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Marquettia americana, 21 Mesostigmata, 60 Nemopteridae, 20 N eoloboqynium americana, 61 Neotropical species of the ant genus Strumigenys group of gundlachi, 37 Neuroptera, 20 Nitidulidae, 35 Olivierina metzeli, 20 Pheidole, 1, 30 Pheidole militicida, 1 Pogonomyrmex, 9, 25 Piesma cinerea, 63 Piesmidae, 63 Release of alarm and attack behav- ior in some New World army ants, 25 Some new synomymy in the Haemo- gamasidae, Laelaptidae and Dip- logyniidae indicated by an exam- ination of Banks’ types of meso- stigmata (Acarina), 60 Spider genus Sty posts, 13 Stephanitis globulifera, 65 Stephanitis pyriodes, 65 Stephanitis rhododendri, 65 Strumigenys , 37 Strumigenys denticulata, 47 Strumigenys eggersi, 46 Strumigenys gundlachi, 40 Strumigenys jamaicensis, 45 Strumigenys subedentata, 48 Strumigenys trieces, 50 Styposis, 13 Styposis a j o, 19 Styposis CHICKERINGI, 15 Styposis cl a u sis, 15 Styposis flavescens , 14 Styposis nicaraguensis, 15 Styposis rancho, 18 Styposis scleropsis, 18 Tamdem running in ants, 29 Theridiidae, 13 Tingidae, 63 William M. Mann, 55 8o PSYCHE A JOURNAL OF ENTOMOLOGY Established in 1874 Vol. 66 March-June, 1959 Nos. 1-2 CONTENTS The Habits of P held 'ole militieida Wheeler (Hymenoptera : Formicidae). W. S. Creighton and M. P. Creighton 1 The Spider Genus Styposis (Araneae: Theridiidae) . H. JV. Levi 13 Fossil Nemopteridae (Neuroptera) . F. M. Carpenter 20 The Release of Alarm and Attack Behavior in Some New World Army Ants. W. L. Brown , Jr 25 Mfl Hi HU1A nK. CAMBRIDGE ENTOMOLOGICAL CLUB Officers for 1958-59 President ....N. Gilham, Harvard University Vice-President M. Parsons, Radcliffe College Secretary A. Stuart, Harvard University Treasurer F. M. Carpenter, Harvard University Executive Committee J. Woodland, Salem State Teachers College S. Duncan, Boston University EDITORIAL BOARD OF PSYCHE F. M. Carpenter (Editor), Professor of Entomology , Harvard U niversity P. J. Darlington, Jr., Head Curator of Recent Insects , Museum of Comparative Zoology W. L. Brown, Jr., Associate Curator of Insects , Museum of Compara- tive Zoology E. 0. Wilson, Associate Professor of Zoology , Harvard University H. Levi, Associate Curator of Arachnology , Museum of Compara- tive Zoology PSYCHE is published quarterly, the issues appearing in March, June, Sep- tember and December. Subscription price, per year, payable in advance: $4.50 to Club members, $5.00 to all other subscribers. Single copies, .$1.25. Checks and remittances should be addressed to Treasurer, Cambridge Ento- mological Club, Biological Laboratories, Harvard University, Cambridge, Mass. Orders for back volumes, missing numbers, notices of change of address, etc., should be sent to the Editorial Office of Psyche, Biological Laboratories, Har- vard University, Cambridge, Mass. IMPORTANT NOTICE TO CONTRIBUTORS Manuscripts intended for publication, books intended for review, and other editorial matter, should be addressed to Professor F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. Authors contributing articles over 8 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 $6.00 per page. The actual cost of preparing cuts for all illustrations must be borne by contributors; the expense for full page plates from line drawings is approximately $8.00 each, and for full page half-tones, $10.00 each; smaller sizes in proportion. AUTHOR’S SEPARATES Reprints of articles may be secured by authors, if they are ordered before, or at the time proofs are received for corrections. The cost of these will be furnished by the Editor on application. The mailing date of the December, 1958 Psyche (Vol. 65, No. 4) will be stated in the next issue. The Lexington Press, Inc., Lexington, Massachusetts PSYCHE Nos. 1-2 Vol. 66 March-June, 1959 THE HABITS OF PHE1DOLE MILITICIDA WHEELER (HYMENOPTERA: FORMICIDAE)1 By Wm. S. Creighton and Martha P. Creighton Department of Biology, City College, New York. During the winter of 1959 the writers were able to study a num- ber of nests of Pheidole militicida Wh. These were situated in an area along the eastern base of the Chiricahua Mountains, six to nine miles north of Portal, Arizona. Extended field studies were made on these colonies and samples taken from them were placed in small Janet nests and observed at the Southwestern Research Station. The need for such work will be clear to anyone familiar with W. M. Wheeler’s remarkable views on the habits of militicida. Wheeler’s initial acquaintance with this ant was made on Novem- ber 10th, 1910, at Benson, Arizona. A day or two later he found other colonies at Hereford. If Wheeler’s hypothesis on the habits of this species is to be appreciated properly, it is important to under- stand exactly what information he secured from these colonies. Be- fore he excavated them, Wheeler found the remains of numerous majors on some of the chaff piles. The material in the Wheeler Col- lection indicates that these remains consisted largely of head capsules which lacked all appendages. If Wheeler had found living majors in the nests that he excavated, these disarticulated remains would have received scant attention. Unfortunately, Wheeler did not find living majors in the colonies which he dug out. But it should be remem- bered that after Wheeler had examined the nests at Benson and Hereford, he knew surprisingly little about the ants that were living in them. He could not even be sure that the majors, whose remains he had discovered on the chaff piles, had come from the nests where he found them. All that Wheeler knew was that he had taken the minor of an undescribed species of Pheidole from these nests and that, after considerable digging, he had been unable to secure any accom- panying majors. Contribution from the Southwestern Research Station of the American Museum of Natural History. I 2 Psyche [March - June The next factual data on militicida came to light a year or two later when W. M, Mann excavated nests near Benson in August and found both majors and minors living in them. These majors corresponded to the remains that Wheeler had taken earlier. With both major and minor castes available, Wheeler was in the position to describe militicida as a new species, which he did in 1915 (1). If Wheeler had been content to let the matter rest with the de- scription of these two castes his action could have been defended not only as sound but also as conservative, for he had waited until the majors and minors had been certainly associated before presenting his description of them. But Wheeler was not content to limit his efforts to the description of militicida. He proceeded to develop a striking hypothesis as to its habits. This was so plausibly presented that one is apt to forget that this highly ingenious account is pri- marily compounded of supposition. Because Wheeler had found no majors in the Benson and Hereford colonies, he assumed that none were present. Because he had found the remains of majors on the chaff piles of these colonies, Wheeler assumed that all the majors in the colonies had been slaughtered by the minors. Having made these two assumptions Wheeler was forced to make others to support them, for this astonishing behavior on the part of the minor clearly demanded explanation. This Wheeler supplied as follows: “It appears, therefore, that all the individuals of this caste (the majors) are regularly killed off by the workers on the approach of winter, probably after they have broken open all the hard seeds col- lected by the workers. Such a slaughter of the members of a large caste during the season when their activities are no longer required, when they would simply be a burden on the colony by consuming stored food and when fresh food cannot be collected, must have great advantages. Although I have never noticed this behavior in any other species of Pheidole , I believe that a study of the harvesting spe- cies with very large-headed soldiers in the deserts of the southwest may bring other similar cases to light.” It is obvious that the most direct way to test Wheeler’s theory would be to examine the nests of militicida during the winter months. If such examinations were properly carried out it would be possible to state with certainty whether or not majors are present in the nests at that time. But subsequent observations on militicida have all been made during the summer. The writers observed this ant at Safford, Arizona, in July 1950. In August, 1952 A. C. Cole studied colonies at Bayard and Deming, New Mexico, and in August 1956 1960] Creighton — Pheidole militicida 3 he found others in the vicinity of Portal, Arizona. Thei information concerning the exact site of the latter colonies, which Dr. Cole obligingly furnished, was a notable help to this study and we wish to express our thanks for it. While the observations just cited added to our knowledge of militicida , none of them could furnish information on winter condi- tions in the nests of this species. Thus when Creighton and Gregg reviewed Wheeler’s hypothesis in 1955 (2), they could do little more than point out the improbability of his explanation. Although it was clear that a number of southwestern species of Pheidole , which have majors with unusually large heads, showed none of the habits predicted by Wheeler, it was still possible that he might have been right in the case of militicida. Positive proof that Wheeler’s account of the habits of militicida is incorrect was secured by the writers in the winter of 1959. During that winter we had fifteen colonies of militicida under field observation on all but a few days. Living majors were taken from thirteen of these colonies during the period between January 8th and March 3rd. Our original attempts to se- cure majors were based upon the obvious method of digging out the colony. This proved to be the worst possible way to get them. Under ordinary conditions only two or three majors stay in the passages near the nest entrance. Since the major of militicida is extraordinarily clumsy, it is seldom able to extricate itself if covered with soil. Hence, it is extremely likely to be missed when the nest is dug out, for the major will often remain perfectly quiet if only a thin layer of soil covers it. To be sure that the majors have not been missed, the soil must be sifted as the nest is excavated. With this in mind it is easy to understand why half the nests which Cole excavated in the summer of 1952 (3) appeared to contain no majors. During the first weeks of this study we faced a similar situation. Three of the eight nests excavated had yielded no majors. The re- maining five had yielded a total of only fourteen majors, of which the greatest number taken from a single nest was five. Our results were, therefore, inconclusive for in no case had numerous majors been found in any nest. Then the junior author hit upon a method of using bait to bring the majors out of the nests. The best bait was found to be various sorts of bread or scrambled eggs. Meat seemed decidedly less at- tractive. 1 he bait was cut into pieces too large for the minors to move and these were placed close to the nest entrance. Phis baiting seldom failed to produce majors in quantity if continued long enough. 4 Psyche [March - June One colony which was baited for five successive days in February yielded a total of seventy-seven majors. Moreover, by using bait majors were secured from nests which had produced none when dug out. For, with one exception, the excavated nests reestablished them- selves after a few weeks. This is clear evidence that these nests had not been fully exposed. The character of the militicida nest would make complete exposure difficult. All the nests that we have encoun- tered have been built in light, friable soil between large stones. As these stones are removed the soil between them crumbles away and this obliterates any passages that were in it. As a result it is usually impossible to follow the passages to any depth and as excavation pro- ceeds there is not the slightest indication of the direction it should take. In our opinion no nest of militicida has yet been completely excavated. In areas where militicida is abundant there are often places where several nest entrances are close together. The distance between the entrances will vary from two to eight feet. It seems impossible at present to state whether each entrance represents a separate nest or whether they all belong to a single elongated nest. The total nest count given in this paper (15) is based upon the latter supposition. The total number of entrances found is five or six times greater. It was soon found that majors and minors from entrances a few feet apart could be mixed together without showing any signs of animosity. At first we accepted this as positive proof that the several entrances all belonged to one nest. But later the disconcerting fact came to light that the same result could be secured with specimens from nests a half a mile or more apart. The only explanation that would seem to fit this surprising behavior is that militicida is almost totally devoid of inter-colony animosity. The situation is made even more inexplicable by the strong animosity which militicida exhibits toward other species of ants. We expected militicida to forage sporadically during the winter but it was a surprise to discover that it is one of the most consistent winter foragers in the area where it occurs. The only other ant which shows comparable activity is Myrmecocystus mimicus Wh. Except for one or two days when rain or snow fell, the minors of militicida were out every day during January, February and early March. As a rule the foraging did not begin until 3 :oo P. M. By that time the surface temperature had reached 6o°F. (i6°C.) or better. Full foraging activity developed when the surface temperature reached 90°F. (3i°C.). During January the surface temperature 1960] Creighton — Pheidolc militicida 5 drops rapidly towards sundown and foraging in that month ordinarily terminates soon after 5 :oo P. M. 'Phis brief period of foraging is extended as the days lengthen and by the end of February the fora- ging lasts about three hours. During the winter months the seeds of two grasses are the principal ones collected. These are the fluff grass, Tridens pulche/lus Hitch and the spike pappus grass, Enneci- pogon desvauxii Beauv.2 Both these grasses fail to lose all their seeds at the end of the growing season, but the number of unshed seeds in the heads is low. Counts on five samples (30 cc. each) of fluff grass heads taken within foraging range of five militicida colonies gave an average of only 4.6 seeds per cubic centimeter. Nevertheless, these residual seeds furnish a steady, if meager, supply for, as the winter advances, the seeds or the spikelets containing them are gradu- ally blown out of the heads and deposited in a thin layer on the surface of the soil wherever there is a windbreak. In the winter months the militicida minors collect their seeds entirely from this layer. During the many days that the foraging minors were observed, not one was seen to ascend a grass stem to get at the seeds. It is difficult to explain this reaction, for the percentage of seeds in the heads is several times as great as that in the layer on the ground. For this layer contains many spikelets that are devoid of seeds and the minors frequently bring these empty envelopes home. On several occasions we took numbers of the spikelets away from the workers as they reached the vicinity of the nest and found that not more than a third of these contained seeds. Despite the short foraging period and the scant seed supply, the militicida minors bring in many seeds, for on warm days foraging is very active. Each nest entrance usually has a single foraging col- umn but sometimes two or three columns may leave the same entrance. The columns are seldom more than fifteen feet long, an indication of the easy availability of the seed supply, regardless of its low yield. Since the rate of seed consumption in the captive colonies was very low, it seems probable that winter foraging augments the number of seeds stored in the nests. The militicida colony is thus provided with an abundant store of seeds which it can use with the arrival of spring. The spring months ’are the driest part of the year in the areas where militicida occurs. At Portal the total precipitation during April, May and June averages 1.52 inches, approximately 8.2% of the an- 2We wish to thank Mr. Joseph Welch, who was working at the Station when this study was made, for his kindness in identifying these and other grasses as well as for helpful suggestions on the terminology of the spikelet. 6 Psyche [March - June nual total of. 18.40 inches (4). In other stations where militicida occurs the percentage of spring precipitation is even lower. As the spring drought begins, the seed supply reaches its lowest ebb, for winter foraging has depleted the meager residue of seeds and no more seeds are likely to be set until after the summer rains. These unfavorable conditions often lead to a suppression of foraging in the spring, but this does not mean that the colony is inactive. For the sexual brood is brought to maturity at this time with the marriage flight following in early July. Thus the heaviest drain on the supply of seeds stored in the nests occurs during the spring months. It is at this time that the stored seeds are broken open and the food in them is made available to the rapidly developing brood. Before taking up the feeding habits of militicida it is advisable to discuss certain features of the grass seeds which they use for food. At maturity a typical grass seed is enclosed in a complicated envelope of bracts. The outermost of these bracts are called glumes, the median ones lemmae and the innermost ones paleae. The lemmae often bear prominent projecting bristles called awns. These parts are attached, close above one another to a much-shortened stem called the rhachilla. Collectively these parts make up a spikelet, which may contain one or more seeds. As a rule when a mature seed is shed, all of the spikelet except the glumes is shed with it. In some grasses this envelope fits the seed tightly, which makes its extraction difficult. In other grasses the envelope is loosely fitted around the seed and its extraction is comparatively easy. The spikelet of E. desvauxii shows the first condition, that of T. pulchellus the second. Since most of the seeds which the minors of militicida bring back to the nests are still in the spikelets, the seeds musts be freed of these envelopes before they can be conveniently stored. Observations on the captive colonies showed that the seeds of T. pulchellus offer no particular problem because of their loose envelopes. The minors have little difficulty in pushing the lemmae aside and can often pull out the seeds without de- taching the surrounding parts. Perhaps this is why the majors so seldom help when pulchellus spikelets are being handled. The spike- lets of E. desvauxii are quite another matter. Each of the close-fitting lemmae has nine slender awns at its tip. These radiate outward at an angle of about 45 degrees and form a complete circle of bristles at the upper end of the spikelet. When attempting to free these seeds of the envelope both majors and minors will grasp the awns in their mandibles. When the major does so the awns are usually broken off, but the minor handles them more gently and when two minors grasp 1960] Creighton — Pheidole militicida 7 the awns at opposite sides of the spikelet and pull in opposite directions they are usually able to break apart the lemmae and thus expose the inner parts of the spikelet. The seed then lies between two paleae but these do not completely enclose it and it is a simple operation for the minors to pick the exposed seed out. When the major extracts the seeds the lower end of the spikelet is grasped in the major’s jaws. Pressure from these either breaks the lemmae apart or, if the seed has exactly the right position as the jaws close, it is forced outward be- tween the lemmae and paleae, which spread enough to let the seed squeeze through. The envelope is often left intact when the seed has been extracted by this method. Whether intact or in fragments, the envelope is then ready to be placed on the chaff pile and the seed extracted from it may be cracked open and eaten or stored in one of the seed chambers. There is abundant evidence that when the seeds are stored they are entire. The minors in our artificial nests spent many hours arranging and rearranging the seeds in groups. We take this to be the equivalent of the packing of the seed chambers in a free nest. No seed in these groups was ever damaged in any way until it was taken out and eaten. Moreover, there were usually a number of partially eaten seeds pres- ent in the nests, for the piilchellus seeds are seldom entirely con- sumed. Such opened seeds invariably shriveled after a day or two and usually moulded a few days later. That cracked or damaged seeds could be stored for months in the seed chambers seems thorough- ly impossible. We were surprised to discover that all the seeds opened in the arti- ficial colonies were opened by the minors. The majors never made the slightest efforts to open the seeds and rarely paid any attention to them once their envelopes had been removed. In an effort to force the majors to crack open seeds, several nests containing only majors were set up. These were liberally supplied with seeds of T. pulchellus. Some of the majors in these nests lived for several weeks but they never made any attempt to open the seeds and ultimately all of them died, apparently from starvation, in the midst of the seeds which could have supplied them with food. The minors open the seeds of pulchellus and desvauxii by gnawing at the pointed end of the seed. Sometimes the seed is held by one minor and gnawed open by another, but a more common method involves only one minor, who places the blunt end of the seed on the floor of the nest and, with the seed held in a vertical position gnaws at its pointed end. The seeds of des- vauxii are entirely consumed but, as noted above, those of pulchellus 8 Psyche [March - June are usually only half eaten. This may be due to the fact that in pulchellus the embryo is confined to the half of the seed which the ants eat, whereas in desvauxii the embryo extends more than three- quarters the length of the seed. As will be seen from the previous paragraphs, the major of militi- cida has a very limited part in the harvesting operations and even this small part can be handled equally well by the minors. It seemed unlikely that the major would be limited to so small a share in the activities of the colony. The first hint that they might perform some unique activity essential to the colony was provided by the majors who came out of the nests during baiting. The junior author, who spent much time at this work, noticed that before the major emerged from the nest it would often stand for a considerable period just in- side the nest entrance. When it did so it was a thorough nuisance to the minors, who had difficulty in getting past what amounted to a road block. When the major finally emerged from the nest it usu- ally opened its jaws to their fullest extent and made short lunges in the direction of small pebbles and bits of grass as though it were trying to bite them. Later it was found that this same lunging and biting response could be elicited by throwing a small beam of light into the opening before the major left it. It was further clear that the primary reason why the major left the nest was not hunger. They almost never went directly to the bait, but wandered around in their clumsy fashion as though they were looking for something else. It was only after considerable patrol that some of them would go to the bait. These responses suggested that the function of the major might be to guard the nest entrance and, to test this, a nest was con- structed which gave them the opportunity to do so. This nest consisted of two chambers connected by a single, long passage which could be blocked or unblocked in the middle without dis- turbing the nest. The block consisted of a cotton plug which could be pushed into the connecting passage through a glass tube set at right angles to it. With the plug in place the nest was divided into two separate chambers; with the plug removed the two chambers communicated with each other through the single connecting passage. After the plug was in place majors and minors of militicida were placed in one chamber and their prospective intruders in the other. The nest was then set aside until both groups were accustomed to their surroundings. Usually it took no more than twelve hours for each group to become thoroughly tranquil and to demonstrate by this tranquility that it was unaware of the other group’s presence nearby. 1960] Creighton — Pheidole militicida 9 The ants selected as intruders were Pogonomyrmex maricopa Wh. and Pogonomyrmex calif ornicus Buck. This choice was made because both species occur in close proximity to militicida colonies in the field and the harvesting activities of all three species lead to frequent encounters outside the nests. The results secured from the experi- mental nest described above were highly interesting. On the removal of the blocking cotton plug both groups would begin to explore the communicating passage. They did so in an entirely different fashion. The Pogonomyrmex workers moved slowly into the passage but rapidly backed out of it when they became aware of the advancing militicida workers. In most cases the militicida minors first entered the passage. Some of them would usually be seized and killed by the Pogonomyrmex workers but others returned to the nest and alerted the majors. When these entered the passage they showed precisely the reactions that they had exhibited around their nest entrances. They advanced very cautiously, with the jaws wide open, and made frequent short lunges in the direction of the Pogonomyrmex Fig. 1 — Major of Pheidole militicida in the defensive posture. workers. As the militicida majors wedged themselves tightly into the passage, three or four ranks deep, the passage was completely blocked and the front face of this block was a highly dangerous area for the Pogonomyrmex workers for it consisted of the closely ap- proximated heads and wide open jaws of the militicida majors. As to what happened next depended on the Pogonomyrmex workers, who would charge up to the barrier and slash at the militicida majors with their mandibles. These attacks were usually futile, for the only exposed parts of the militicida major which could be damaged were IO Psyche [March - June the antennae and these were held so closely against the head that the Pogonomyrmex workers were seldom able to grasp them. If these attacks were vigorously pressed the militicida major usually stood perfectly still and waited until the mandibles of its opponent were near its own. It then lunged forward, closed its jaws on the mandible of the Pogonomyrmex worker and attempted to break of! the crushed mandible. The majors did not always succeed in doing so, particularly in the case of maricopa, whose heavy mandibles are hard to break, but they seldom failed to mangle the mandible so badly that it was useless. It may be added that this attack on the mandible is deliberate, for the militicida major will rarely strike at other parts when these are presented. We have repeatedly seen the Pogonomyrmex workers thrust their antennae or legs between the open jaws of the militicida major without causing the major to strike. They do not do so until there is a good chance that the mandible can be grasped and they rarely miss their target. After a number of Pogonomyrmex workers had been put out of action with useless mandibles, or sooner if the Pogonomyrmex workers did not press the attack vigorously, the militicida majors emerged from the passage and began a different sort of action. They no longer faced their opponents and struck at their mandibles but approached them from the rear and struck at the thorax or the petiolar nodes. As a result, most of the Pogonomyrmex workers were ultimately cut in two, either at the petiole or behind the pronotum. In this more open fighting it was also obvious that the petiolar nodes and the mesothoracic area were the principal targets. An examination of the Pogonomyr- mex workers at the end of an engagement always showed much damage to mandibles, thorax and petiolar nodes and surprisingly little damage to legs and antennae. In short, there is nothing hap- hazard about the way in which the militicida majors deal with their opponents; they only strike at parts which will put their opponents out of action or kill them. It is clear that their method is highly effective for it was only occasionally that the Pogonomyrmex workers got the better of the engagement. Even when they outnumbered the militicida majors they often failed to kill a single one of them and when they did so it was usually a result of the militicida major having been stung. This incapacitates them but does not immediatelv kill them. It should be clear that the activities just described are considera- bly more methodical that the ordinary scrimmages between fighting ants. In these activities the militicida major shows an efficiency that 1960] Creighton — Phcidole militicida 1 1 is completely unlike its bumbling efforts elsewhere. This, plus the fact that these responses are repeated with surprising exactness time after time, and by majors from different nests, leads us to conclude that they are the normal guarding responses of the militicida major. If this is true the major of militicida is best regarded as a soldier. Its role in the harvesting activities of the colony is slight and it is not primarily a seed-crusher, as has been mistakenly supposed. The defensive activities of the militicida major probably account for the mutilated remains which Wheeler found when he discovered this species, for the defending majors do not always dispatch the in- truders without loss to themselves. This seems a more probable ex- planation than that proposed by Creighton and Gregg in 1955 (2), who suggested that the accumulation of dead majors discovered by Wheeler might have been a result of the high death rate of that caste during the peak of the harvest season. This now seems unlikely, for the death rate of the majors during the winter months has proved to be extremely low. Most of the fifteen colonies that we studied dis- carded no dead majors during the three months that they were under observation. Only four dead majors were placed on the chaff piles during this period. Each of these was carefully examined under a binocular microscope for signs of mutilation and each was found to be completely undamaged with not even a tarsal claw missing. This disposes of the last bit of evidence on which Wheeler based his hypothesis, for it is now clear that the minors of militicida neither kill the majors nor cut them to pieces after they have died. Thus Wheeler’s views of the habits of militicida have proved to be at total variance with what these habits actually are. In conclusion, it may be noted that militicida is a very difficult subject for investigations in artificial nests. This species is unusually sensitive both to temperature and humidity. Temperatures below freezing invariably kill the ants and they are almost as seriously affected by a heavy condensation of water on the glass of the nests. I hey seem unable to keep out of the water droplets and many of the minors die in them. They also die if the nest is too dry. This sensitivity made it difficult to maintain the captive colonies for any length of time. At the suggestion of Dr. Robert Chew, of the De- partment of Biology of the University of Southern California, the artificial nests were placed on a rack, in a covered aquarium, above a saturated solution of sodium chloride. In a closed system this should maintain a constant humidity of 78% of saturation regardless of temperature. The arrangement proved eminently satisfactory. It 12 Psyche [March - June not only reduced condensation but also largely eliminated the need for adding water to the individual nests. We wish to thank Dr, Chew for suggesting a method which not only benefited this study but which should be of value to anyone faced with the maintainance of a “fussy” species in the laboratory. Literature Cited. 1. Wheeler, W. M., 1915. Bull. Amer. Mus. Nat. Hist. 34: 389-421. 2. Creighton, W. S. and R.E. Gregg, 1955. Univ. Colorado Studies, Series in Biol. No. 3: 1953. 1-46. 3. Cole, A. C., 1953. Jour. Tenn. Acad. Sci. 28: 297-298. 4. 19+1. Climate and man. U. S. Department of Agriculture. Yearbook. Part 5. THE SPIDER GENUS STYPOSIS (ARANEAE, THERIDIIDAE) 1 By Herbert W. Levi Museum of Comparative Zoology, Harvard University Theridiid spiders usually have eight eyes. Only recently the genus Archerius Levi (1957, Trans. Amer. Micros. Soc., 76: 114) was described from some California specimens having only six eyes. Since then an eight-eyed species from Japan was seen, a species having the anterior median eyes small. The number of eyes, then, may not be of generic importance. Since the work on Archerius , another theridiid genus has been found in which the eye number is variable. In some species the anterior median eyes are present but small ; other species lack them entirely. Judging by their similar appearance and by the genital structure, the species are closely related and belong to the genus Styposis. A grant from the National Institutes of Health (E-1944) and a National Science Foundation grant (G-4317) made the completion of this paper possible. I would like to thank Dr. W. J. Gertsch and Dr. A. M. Chickering for specimens used and Professor M. Vachon for permitting me to examine S. flavescens in the Museum National d’Histoire Naturelle in Paris. Styposis Simon Styposis Simon, 1894, Histoire Naturelle des Araignees, 1: 592. Type species by original designation and monotypy: S. flavescens Simon. Small unpigmented spiders, less than 2 mm. total length. In some species carapace almost circular, weakly sclerotized, with six large eyes (except all eyes relatively small in S. ajo, Figure 27) arranged in two groups of three eyes touching each other (Figs. 1, 5, 10, 11). In other species ( S . flavescens , S. scleropsis) carapace longer than wide, sclerotized, with raised reticulate pattern (Figs. 18, 24). In all spe- cies anterior median eyes absent or minute, their maximum diameter equal to radius of posterior medians. Chelicerae weak with one or two teeth on anterior margin (Fig 11), probably none on posterior. Legs fairly long; in some species patella and tibia 2.7 times carapace length (S. clausis) , in others shorter 1.4 times carapace length (S. ajo , S. flavescens) . Legs with many setae; comb on fourth tarsus greatly reduced, its setae almost smooth. Abdomen soft, sometimes wider than long, or high. Colulus two minute hairs slightly anterior of Tublished with the aid of a grant from the Museum of Comparative Zoology at Harvard College. 13 14 Psyche [March - June usual position, difficult to discern and could not be found in some species. Female genitalia with two seminal receptacles. Male palpus weak- ly sclerotized, parts translucent and difficult to delineate. Bulb twisted so that embolus faces the outside (and is partly hidden by the' cym- bium), and median apophysis faces ventrally (Figs. 6, 9, 16, 19). Median apophysis (M in Figs. 8, 14, 15, 20) a very large structure, functional, its end lying against the paracymbial hook (Fig. 14) at the distal end of the cymbium. Radix (R) present. The large struc- ture supporting the embolus in S. chickeringi and S. flavescens prob- ably is the radix (R in Figs. 14, 15, 20) . Six species are known from Central America and northern South America and one from the southwestern United States. Styposis differs from Archerius by lacking the large colulus and by being less sclerotized. It differs from Sphyrotinus by having the anter- ior median eyes reduced in size or absent and by the unusual position of the bulb in the palpal cymbium. Styposis flavescens Simon Figures 7, 18-22 Styposis flavescens Simon, 1894, Histoire Naturelle des Araignees, 1:592. Fig. 599. 9 type from Venezuela, in the Museum National d’Histoire Natur- elle, Paris, examined. Description. Carapace, sternum brown. Legs light brown. Abdo- men whitish, without pigment. Carapace longer than wide. Cephalo- thorax heavily sclerotized ; carapace and sternum with a net-like sclerotized pattern (Fig. 18). Diameter of anterior median eyes about equal to the radius of posterior medians. Anterior median e3^es their radius apart, their radius from laterals. Posterior median eyes one-third diameter apart, two-thirds from laterals. Abdomen suboval in shape. Total length of female 1.4 mm. Carapace, 0.68 mm. long, 0.53 mm wide. First femur, 0.89 mm.; patella and tibia, 0.92 mm.; metatarsus 0.62 mm; tarsus, 0.41 mm. Second patella and tibia, 0.68 mm.; third, 0.47 mm.; fourth, 0.68 mm. Total length of male 1.5 mm. Carapace 0.75 mm. long, 0.56 mm. wide. First femur, 1.04 mm.; patella and tibia, 1.22 mm.; metatarsus, 0.81 mm.; tarsus, 0.47 mm. Second patella and tibia, 0.83 mm.; third, 0.59 mm.; fourth, 0.70 mm. The epigynum (Figs. 7, 22) has a slight median projection. The palpus (Figs. 19, 20) has a median apophysis with two projecting prongs, easily seen in lateral view. (Fig. 7 was made from the type, Figs. 18-22 from specimens collected in Nicaragua). 1960] Levi Spider genus Styposis 15 Record. Nicaragua. Musawas, Waspuc River, Oct. 10-31, 1955, 9, cT, Nov. 1-4, 1955, ?, cT (B. Malkin). Styposis chickeringi, new species Figures 10-17 Types. Male holotype and female paratypes from El Valle, Pana- ma, July 1936 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Spider colorless, except for silvery eyes and spots of pigment where anterior median eyes usually are. Posterior median eyes one diameter apart, touching laterals. Chelicerae with one blunt tooth (Fig. 11). Abdomen wider than long (Fig. 10). Measure- ments of male holotype, total length 1.3 mm. Carapace, 0.77 mm. long, 0.72 mm. wide. First femur, 1.50 mm.; patella and tibia, 1.82 mm.; metatarsus, 1.32 mm.; tarsus, 0.44 mm. Second patella and tibia, 1.22 mm.; third, 0.75 mm.; fourth, 0.96 mm. Measurements of female paratype, total length 1.3 mm. Carapace, 0.60 mm. long, 0.55 mm. wide. First femur, 1.25 mm.; patella and tibia, 1.35 mm.; metatarsus, 0.96 mm. ; tarsus, 0.36 mm. Second patella and tibia, 0.88 mm. ; third, 0.52 mm. ; fourth, 0.82 mm. Diagnosis. The genitalia (Figs. 12, 13, 16, 17) as well as the eye distance separate this species from the others of this genus. Styposis nicaraguensis, new species Figures 8-9 Type. Male type from Musawas, Waspuc River, Nicaragua, Oc- tober 10, 31, 1955 (B. Malkin), in the American Museum of Natural History. Description. Spider slightly yellowish. Anterior median eyes ab- sent. Anterior eyes slightly more than their diameter apart. Posterior median eyes their radius apart, touching laterals. Chelicerae with two teeth on the anterior margin. Total length of male type 1.4 mm. Carapace 0.62 mm. long, 0.55 mm. wide. First femur, 1.46 mm.; patella and tibia, 1.55 mm.; metatarsus 1.03 mm.; tarsus, 0.47 mm. Second patella and tibia, 1.0 mm.; third, 0.56 mm.; fourth, 0.94 mm. Diagnosis. The male palpus (Fig. 9) distinguishes this species. Styposis clausis, new species Figure 1-4 Types. Female holotype, female paratype from Forest Reserve, Canal Zone, July 4-6, 1939 (A. M. Chickering), in the Museum of Comparative Zoology. i6 Figs. 1-4. Styposis clausis, new species. 1. Female. 2. Female genitalia, dorsal view. 3. Epigynum. 4. Epigynum from side. Figs. 5-6. S. rancho, new species. 5. Carapace of male. 6. Left palpus. Fig. 7. S. flavcscens Simon, epigynum. Figs. 8-9. S. nicaraguensis, new species, palpus. 8. Expanded, ectal view. 9. Ventral view. Figs. 10-17. S. chickeringi, new species. 10. Female. 11. Head of female. 12. Female genitalia, dorsal view. 13. Epigynum. 14. Palpus expanded, subventral view. 15. Palpus expanded, ectal view. 16. Palpus of type. 17. Right palpus of paratype drawn at slightly different angle. Figs. 18-22. S. flavescens Simon. 18. Carapace of female. 19-20. Left pal- pus. 19. Ventral view. 20. Ectal view, expanded. 21. Female genitalia, dorsal view. 22. Epigynum. Figs. 23-24. S. scleropsis, new species. 23. Pal- pus. 24. Carapace of male. Figs. 25-27. S. ajo , new species. 25. Female gen- italia, dorsal view. 26. Epigynum. 27. Carapace of female. (Abbreviations: C, conductor; E, embolus; M, median apophysis; R, radix; T, tegulum). Description. Carapace slightly yellowish, otherwise colorless. Six eyes. Anterior eyes one diameter apart. Posterior median eyes one- third diameter apart, almost touching laterals. Chelicerae with one blunt tooth on anterior margin. Abdomen wider than long when filled with eggs, otherwise longer than wide. It is not known whether the structure on the epigynum (Figs. 3, 4) is an epigynal plug or part of the epigynum. As the structures of the two specimens examined differ slightly, it probably is an epigynal plug. Total length 1.7 mm. Cara- pace, 0.73 mm. long, 0.69 mm. wide. First femur, 1.72 mm.; patella and tibia, 1.95 mm.; metatarsus, 1.33 mm.; tarsus, 0.52 mm. Second patella and tibia, 1.14 mm.; third, 0.73 mm.; fourth, 1.06 mm. Diagnosis. The arrangement of the eyes (Fig. 1) and genitalia (Fig. 2) separate this species from others. i8 Psyche [March - June Styposis rancho, new species Figures 5-6 Type. Male type from Rancho Grande, Venezuela, December 20, 1954 (A. M. Nadler), in the American Museum of Natural History. Description. Carapace with a slightly dusky border. Abdomen with some white pigment on dorsum. Anterior median eyes minute. Anterior lateral eyes about one diameter apart. Posterior median eyes one-third of their diameter apart. Chelicerae with one tooth on the anterior margin. Abdomen probably wider than long, damaged in the type specimen. Total length 1.6 mm. Carapace 0.62 mm. long, 0.54 mm. wide. First femur, 0.96 mm.; patella and tibia, 1.02 mm.; metatarsus, 0.75 mm.; tarsus, 0.34 mm. Second patella and tibia, 0.77 mm.; third, 0.45 mm.; fourth, 0.65 mm. Diagnosis. The eye arrangement (Fig. 5) and the palpus (Fig. 6) separate this species from others. Styposis scleropsis, new species Figures 23, 24 Type. Male from Summit, Panama Canal Zone, August 23-28, 1950 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, sternum, legs light brown. Abdomen gray- ish white. Carapace sclerotized, reticulate. Cephalothorax slightly elongated behind (Fig. 24). Anterior median eyes half the diameter of other eyes. Anterior median eyes their radius apart, two-thirds diameter from laterals. Posterior median eyes almost touching, their radius from laterals. Chelicerae heavy with a large tooth far removed from base of fang and a smaller tooth on base of larger one. Abdo- men higher than long. Around the spinnerets a very lightly sclero- tized ring, hardly different in color from less sclerotized portions. Colulus with two setae quite far anterior. Total length 1.2 mm. Carapace, 0.78 mm. long, 0.52 mm. wide. First femur, 1.00 mm.; patella and tibia, 1.18 mm.; metatarsus, 0.65 mm.; tarsus, 0.39 mm. Second patella and tibia, 0.76 mm. ; third, 0.48 mm. ; fourth, 0.66 mm. Diagnosis. This species resembles Cerocida in the slightly elong- ated cephalothorax and the high abdomen. However, the small anter- ior median eyes and the palpal structure place it in Styposis. The shape of the carapace and the transparency of the palpal sclerites (Fig. 23) separate it from other species. 1960] Levi — Spider genus Styposis 19 Styposis ajo, new species Figures 25-27 Type. Female from 30 miles south of Ajo, Pima County, Arizona, January 4, 1941 (S. and D. Mulaik), in the American Museum of Natural History. Description. Colorless except for pigment around eyes. Diameter of anterior median eyes less than half that of others. Anterior median eyes two and one-half diameters apart, one diameter from laterals. Posterior median eyes one diameter apart, their radius from laterals. Abdomen oval, longer than wide. Total length 1.6 mm. Carapace, 0.60 mm. long, 0.55 mm. wide. First femur, 0.78 mm.; patella and tibia, 0.88 mm. ; metatarsus, 0.62 mm. ; tarsus, 0.39 mm. Second patella and tibia, 0.67 mm.; third, 0.52 mm.; fourth, 0.83 mm. Diagnosis. This species differs from others by the shape of the epigynum (Fig. 26). It is the only member of the genus from north of Central America. FOSSIL NEMOPTERIDAE (NEUROPTERA).1 By F. M. Carpenter Harvard University The living members of the neuropterous family Nemopteridae have a wide but irregular geographical distribution. Although they occur in southern Europe, Asia Minor, India, South America and Africa, they have not been found in North America. However, a fossil species, Halter americana, was described by Cockerell in 1907 from the Florissant shales in Colorado.2 The published description of this fossil, unfortunately, was too brief and incomplete to be of use to specialists on Nemopteridae. Navas, who subsequently examined the type specimen in the British Museum (Natural History), pub- lished a new, but very inadequate, account of the fossil in 1913, erected a new genus, Marquettia , for it, and included a rough figure of the fore wing. The present paper is a more detailed description of the specimen, made in connection with my preparation of the insect part of the Treatise on Invertebrate Palaeontology. I examined the type of americana in the British Museum in 1938, and Dr. R. Baker, of the Department of Palaeontology of the Mu- seum, has recently sent me a series of excellent photographs of it. A second specimen of americana , contained in the Natural History Mu- seum of the University of Colorado, has been loaned to me for study by the director of the Museum, Dr. Hugo Rodeck; it has added some significant features to our knowledge of the species. I have taken this occasion to discuss briefly the nemopterid Olivierina metzeli, which Pierce and Kirkby have recently described from an Oligocene deposit in Montana. Unfortunately, the generic classification of the living Nemopteridae is far from satisfactory. Navas’ two revisional studies were published nearly fifty years ago (1910, 1912), and the generic classification in- cluded there was mainly an arbitrary one. The genera were based almost entirely upon the shapes of the hind wings, without regard to the probably occurrence of convergence in several lines of evolution. The venation of the fore wing is surprisingly constant throughout the entire group with the exception of a very few species in which the Tublished with the aid of a grant from the Museum of Comparative Zoology at Harvard College. This fossil naturally aroused much interest. It was selected by the editors of the Entomological News (19:34, 1908) as the most remarkable insect made known during the year 1907, and it was reproduced on the cover of the issues of the News for 1908. 20 1960] Carpenter — Fossil N emopteridae 21 pattern is much reduced. A satisfactory generic classification of the nemopterids will almost certainly not be achieved until the terminal abdominal segments of the males have been studied in detail for the majority of the species described from all parts of the world. The generic assignments of the fossils discussed below must, of course, be made mainly on the basis of the venation of the fore wing and the general shape of the hind wing, these being the characteristics which have been used in the classification of the living species up to the present time. Genus Marquettia Navas Navas, 1913, Mem. Real Acad. Cien. y Artes de Barcelona, 10:7 Head shaped much as in Lertha, the rostrum present but not elong- ate. Front wing broadly oval, with a venational pattern more or less characteristic of that found in the tribe Stenonemiini (Orfila, 1954). The radial sector is extensively branched, having ten main branches in addition to the anterior media.3 The pterostigma is small and it was probably very light in color in the living insect, as in Lertha . The hind wing is slender, about the same as in Kirbynict in general form, except that it has a more rounded apex, like that of Olivierina and Halterina; it has two dilations, the dilated portion being nearly uni- formly dark in color. Type species: Halter americana Cockerell. As pointed out above, this genus is related, so far as we can see from the known structure, to several stenonemiine genera, but it stands apart from all of these by the extensive development of the radial sector. In this respect, I consider the genus Marquettia to be the most primitive of the genera of the Nemopteridae known at the present time. Marquettia americana (Cockerell) Plate 1 and text figure 1. Halter americana Cockerell, 1907, Science, 26:466 Halter americana Cockerell, 1908, Pop. Sci. Mo., 72: 125 Marquettia americana Navas, 1913, Mem. Real Acad. Cien. y Artes de Barcelona, 10:484, fig. 4. Fore wing, length, 31 mm.; width, 10 mm. Length of hind wing, 44 mm.; length of body, 16 mm.; length of beak, 2.5 mm. The vena- 3I have used here the venational nomenclature proposed by Dr. P. A. Adams in his “Studies in the Neuroptera, with Special Reference to Wing Structure and Evolution in the Osmyloidea” (unpublished doctoral thesis, Harvard Uni- versity, June 1958). 22 Psyche [March - June tion is typical of that of the tribe Stenonemiini, as shown in text figure i, with the exception of the presence of the numerous branches of the radial sector. The hind wing, which is about one and one-half times the length of the fore wing, has a slender petiole and terminates in two conspicuous dilations. The distal dilation appears to be at the termination of the wing; there does not seem to be a terminal exten- sion, tapering more nearly to a point, as Navas indicated by dotted lines in his figure 4 (1913). The two dilations are almost certainly somewhat twisted as the fossil is preserved ; this, of course, is what one would expect from the condition of the hind wing as it occurs in most living specimens of the family. The petiole of the wing is hyaline, but the two dilations are almost solidly dark in color, although there is an indication of a small hyaline area in the region of the twisting; this light area could, of course, be due to peculiarities of preservation, although the two light areas are in the same position in both of the hind wings. Holotype: This consists of a nearly complete specimen with all four wings outstretched (plate 1), with the venation of the front wings very clear ; collected in the Florissant shales, Colorado ;4 deposited in the Department of Palaeontology, British Museum (Natural His- tory). The species which Pierce and Kirkby described (1959) from Oli- gocene shales in Montana as Olivierina metzeli is apparently closely related to americana and may in fact be that species. The size of the insect is virtually identical with that of americana. Unfortunately, the specimen on which the metzeli is based is poorly preserved; the venation of the fore wing is not discernible, and the form of the hind wing is not clearly shown. The authors state that the dilated part of the hind wing narrows for a short distance and widens again to a narrow inflation. Mrs. Kirkby informs me that, though the apical portion is not complete in the type specimen, there is enough evidence preserved to show that the “paddle” widens again. Pierce and Kirkby apparently incorrectly interpreted Cockerell’s description of his americana and they were unaware of Navas’ published redescrip- tion of americana and of his establishment of the genus Marquettia. In his account of americana , Cockerell stated that the black area of 4Formerly thought to be of Miocene age but now usually considered Oligo- cene. Explanation of Plate 1 Marquettia americana (Cock.). Photograph of holotype [Courtesy of British Museum (Natural History)]. Psyche, 1959 Vol. 66, Plate 1 Carpenter — Fossil Nemopteridae 24 Psyche [March - June the hind wing was solid and continuous, not broken into two as in the living Halter extensa. That he was referring to the interruption of the black area and not to the “paddle” is shown by his previous statement that the hind wing had an apical fiddle-shaped expansion, which was dark-colored. Pierce and Kirkby apparently concluded that americana had a single, undivided dilation. The two dilations are shown in Navas’ figure of the type ( 1913) . Since metzeli has two dilations of the hind wing, as in americana , and resembles that species in all other known features, it may most reasonably be placed in the genus Marquettia , to which it is hereby assigned. A generic revision of all known nemopterids may result in consid- erably generic synonymy, with Marquettia included. However, until such a revision is made, I believe Marquettia should be retained for these two fossil Nearctic species. Text figure 1. Fore and hind wings of Marquettia americana (Cock.). Drawing based mainly on holotype, with additional details of specimen no. 4514, University of Colorado Museum. References Cockerell, T. D. A., 1907, Some Old World Types of Insects in the Miocene of Colorado, Science, 26: 446-447. 1908, Florissant: A Miocene Pompeii, Pop. Sci. Mo., 73: 124-125. Navas, R. P. L., 1910. Monografia de los Nemopteridos, Mem. Real Acad. Cien. y Artes de Barcelona, 8: 341-408. 1912, Family Nemopteridae, Neuroptera, Genera Insec- torum, 136: 1-23. 1913, Mis Excursiones por el Extranjero, Mem. Real Acad. Cien. y Artes, 10: 7-9, fig. 4. Orfila, R. N., 1954, Un Nuevo “Nemopteridae” Americano, Rev. Soc. Ent. Argentina, 17:29-32. Pierce, W. Dwight and Ruth A. Kirkby, 1959, Fossil Insects from Montana: A New Fossil Nemopterid (Neuroptera), Bull. So. Calif. Acad. Sci., 58: 47-50. THE RELEASE OF ALARM AND ATTACK BEHAVIOR IN SOME NEW WORLD ARMY ANTS By William L. Brown, Jr. Museum of Comparative Zoology Harvard University The effect of odoriferous mandibular gland secretions (phero- mones) as releasers of alarm behavior in Pogonomyrmex and some other ants has recently been investigated by Wilson (1959 and pers. commun.). It is not known how widely among the subfamilies of ants that the same or an homologous effect may occur, but Wilson, myself and others have been able to detect various pungent odors originating from the heads of worker ants in the subfamilies Myr- micinae ( Pogonomyrmex , Solenopsis) , Ponerinae (Paraponera, Meso- ponera , Leptogenys) , and Formicinae (Acanthomyops, Lasius). That these substances so often originate in the head is suggestive of a similar source and behavioral function, but in most cases experimental evidence of their function is lacking.1 A recent study trip to the Smithsonian Institution’s Canal Zone Biological Area station on Barro Colorado Island" afforded a chance to study alarm behavior in the New World army ants (Dorylinae, tribe Ecitonini). Since the trip had other primary purposes, the army ant observations were rather limited, and some obvious tests that suggest themselves, such as trials of interspecific reactions, were not carried out. The results obtained, however, do indicate that the detailed study of army ant pheromones would be a rewarding one. Schneirla (1956 and earlier papers) has made intensive studies of the army ants of Barro Colorado, and has outlined many aspects of their biology, including the remarkably regular nomadic habits of the Eciton species. The first species I studied, E. hamatum (Fabri- cius), was also the one given the most attention by Schneirla and his colleagues; it is the most conspicuous army ant on the island. A trunk trail of E. hamatum was found at midday running along the edge of a buttress root of a forest tree. The two-way stream of ants varied from two to six individuals in width, and the incoming ants bore much booty, mostly the pupae of the fungus-growing ant, 3I have found that ants of the Australian formicine genus Calomyrmex produce a bright yellow or scarlet droplet at the base of each mandible when roughly disturbed, but it is not known whether this substance is odori- ferous. The trip was supported by a grant from the Milton Fund of Harvard University. 25 26 Psyche [March - June Atta. A faint “meaty” odor was noticed as I brought my face close to the column ; this odor increased markedly when I agitated a part of the column with my machete, and the ants savagely attacked the blade, biting and attempting to sting. (The meatlike odor of some New World army ants has been well known for years.) Selecting one of the large soldiers, with its conspicuous, shiny whitish head and long hooked jaws, I removed the head with forceps and placed it in the column, where it was at once vigorously attacked, bitten and stung by a mass of workers and soldiers that collected around it. The actively struggling body of this same soldier was then placed in the column at another point; the headless body, despite its consider- able and highly irregular activity, attracted only momentary antennal play from passing nestmates. Several more trials like this were made at different points along the column ; it was found that soldier heads attracted more attackers than did the much smaller worker heads tried, and also that freshly crushed soldier heads drew a stronger attack than intact ones. To my nose, the strength of the meaty odor in each case appeared to be roughly proportional to the number and activity of the attackers, and I could detect none of this odor from the postcephalic part of the body after the heads had been removed. (Later, in the laboratory, crushed soldier heads of E. hamatum were sniffed by two colleagues and myself, and compared directly with the odor of beef and vege- table bouillon cubes, with the result that all of us found the odors close if not identical.) Next, small dead twigs were broken in half, and one half rubbed against the crushed head of a worker. Such twigs when placed in the column were subjected to an attack as massive and vigorous as were the crushed heads alone. The untreated half of each twig, serving as a control placed at another point along the column, never re- ceived more than fleeting attention from passing soldiers and workers. Similar results were consistently obtained with another column of E . hamatum found on a subsequent day. The same kind of investigation was carried out briefly with a column of Nomamyrmex esenbecki (Westwood) found in the forest, utilizing the large and medium-sized workers for the decapitation test (this species lacks large soldiers). To my nose, crushed heads of this species have a different and weaker odor than do E. hajnatum workers of the same size, but the alarm and attack reaction they pro- voked was similar to that of hamatum in intensity. The test was also made with soldiers from a raiding swarm of 1960] Brown — A rmy Ants 27 Labidus praedator (Fr. Smith) found in the Laboratory Clearing on Barro Colorado, and results were again similar to those of the first trials with E. hamatum, I could not be sure in this case that I detected any odor at all from the crushed soldier heads of L. praedator , but the intensity of attack on these heads indicated that an odor must have been readily detectable by the workers participating. In summary, the detached heads of workers or soldiers of these army ants from three different genera were attacked by their nest- mates when placed in the foraging column, while the remainder of the body evoked little or no response. Objects smeared with the substance from the crushed heads of soldiers drew intense attack, while untreated control objects did not. It seems likely that a phero- mone originating in the head, and most likely specifically in the mandibular glands, occurs in the army ants as well as in Pogonomyr- mex, and that its function is similar. Apparently, the ants will attack any strange object presented close to the center of concentration of the alarm odor. The question arises as to how the individuals that are disturbed into producing the alarm odor are not themselves attacked in the normal course of events. Two of the most likely answers that sug- gest themselves to me are: (1) workers and soldiers, when normally disturbed, give off much lesser amounts of the alarm odor than is produced when a head is severed from the body or crushed, and the attack on severed heads or crushed heads represents a supernormal stimulus; (2) the postcephalic part of the ant’s body bears some quality, probably again pheromonal in nature, that neutralizes attack behavior that might otherwise be released in nestmates. The second answer is favored as an hypothesis in the absence of further experi- ments. If a given soldier were protected by such a neutralizer or “identification pheromone,” or by a “nest odor,” it would presum- ably be able to indicate by means of alarm odor release the locality of an intruding disturbance without itself being subject to attack by its nestmates. I should like to acknowledge helpful suggestions made by Prof. E. O. Wilson during the course of the work and the preparation of the manuscript. References Schneirla, T. C. 1956. The army ants. In Report of the Smithsonian Institution for 1955, pp. 379-406 (with bibliography). Wilson, E.O. 1959. (1958). A chemical releaser of alarm and digging behavior in the ant Pogonomyrtnex badius Latreille). Psyche. 65: 41-51. CAMBRIDGE ENTOMOLOGICAL CLUB A regular meeting of the Club is held on the second Tuesday of each month October through May at 8:00 p. m. in Room B-455, Biological Laboratories, Divinity Ave., Cambridge. Entomologists visiting Boston are cordially invited to attend. BACK VOLUMES OF PSYCHE The Cambridge Entomological Club is able to offer for sale the following volumes of Psyche. Volumes 3, 4, 5, 6, 7, 8, each covering a period of three years, $8.00 each. Volumes 10, 14, 17 to 26, each covering a single year, $2.00 each. Volumes 27 to 53, each covering a single year, $2.50. Volumes 54 to 65, each covering a single year, $3.00. Some other volumes, lacking certain issues, are also available (information upon request). Orders for 2 or more volumes subject to a discount of 10%. Orders for 10 or more volumes subject to a discount of 20%. All orders should be addressed to F. M. Carpenter, Editor of Psyche, Biological Laboratories, Harvard University, Cambridge, Mass. 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 38, Mass. r\ s 6? 0.48 mm 3, Mandibles longer and slender (MI 56 or more) ; smaller species, worker HL mostly 0.48 mm. or less 4. 3. Mandible very short in proportion to head (MI 48 in unique holotype), with exactly three small preapical teeth; ground pilos- ity of head nearly or quite obsolete; pronotum markedly flattened (Fig. 8; Costa Rica) trieces Brown Mandible a little longer proportionate to head (MI 53-54), with more than three preapical denticles ; ground pilosity abundant and conspicuous on head ; promesonotum strongly rounded, not de- pressed (Fig. 9; s. Mexico to s. Brazil) subedentata Mayr 4. Mandibles very long and slender (MI > 70), bowed outward (Fig. 3; Trinidad to n. Argentina) denticulata Mayr Mandibles not so long (MI < 70), their shafts approximately straight 5. 5. Spongiform appendages small but distinctly developed (Fig. 5) ; gastric dorsum of worker (when clean) predominantly smooth and shining, with a few basal costulae; female gaster commonly shagreened (Fig. 1 ; Caribbean countries) gundlachi (Roger) Spongiform appendages obsolete (Fig. 6) ; gastric dorsum with distinct fine, mostly opaque reticulation (Fig. 2; Brazil, Bolivia, Caribbean countries) eggersi Emery 52 Psyche [September References Brown, W. L., Jr. 1948. A preliminary revision of the higher Dacetini. Trans. Amer. Ent. Soc., Philadelphia, 74: 101-129. Emery, C. 1890. Studi sulle formiche della faune neotropica TV. Bull. Soc. Ent. Ital., 22: 38-80, pi. 5-9. Kempf, W. W. 1958. The ants of the tribe Dacetini in the State of Sa~b Paulo, Brazil, with the description of a new species of Strumig enys. Stud. Ent. (n. s.), Petropolis, Brazil, 1: 553-560. Mayr, G. 1887. Sudamerikanische Formiciden. Verh. zool.-bot. Ges. Wien, 37: 511-632. Roger, J. 1862. Einige neue exotische Ameisen-Gattungen und Arten. Ber- lin. ent. Zeitschr., 6: 233-262, 1. pi. 1863. Verzeichniss der Formieiden-Gattungen und Arten. Berlin. Santschi, F. 1930. Quelques fourmis de Cuba et du Bresil. Bull. Soc. R. Ent. Egypte (n. s.), pp. 75-83. 1931. Fourmis de Cuba et de Panama. Rev. Ent., Rio de Janeiro, 1:265-282. Wasmann, E. 1915. Das Gesellschaftsleben der Ameisen, I; as cited in Wheeler, W. M., 1936, Proc. Amer. Acad. Arts Sci., 71: 228-229. Weber, N. A. 1934. Notes on neotropical ants, including the descriptions of new forms. Rev. Ent., Rio de Janeiro, 4: 22-59. 1952. Biological notes on Dacetini. Amer. Mus. Novit., 1554: 1-7. CAMBRIDGE ENTOMOLOGICAL CLUB A regular meeting of the Club is held on the second Tuesday of each month October through May at 8:00 p. m. in Room B-455, Biological Laboratories, Divinity Ave., Cambridge. Entomologists visiting Boston are cordially invited to attend. BACK VOLUMES OF PSYCHE The Cambridge Entomological Club is able to offer for sale the following volumes of Psyche. Volumes 3, 4, 5, 6, 7, 8, each covering a period of three years, $8.00 each. Volumes 10, 14, 17 to 26, each covering a single year, $2.00 each. Volumes 27 to 53, each covering a single year, $2.50. Volumes 54 to 65, each covering a single year, $3.00. Some other volumes, lacking certain issues, are also available (information upon request). Orders for 2 or more volumes subject to a discount of 10%. Orders for 10 or more volumes subject to a discount of 20%. All orders should be addressed to F. M. Carpenter, Editor of Psyche, Biological Laboratories, Harvard University, Cambridge, Mass. 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 38, Mass. PSYCHE A JOURNAL OF ENTOMOLOGY Established in 1874 Vol. 66 December, 1959 No. 4 CONTENTS William M. Mann. E. O. Wilson 55 Some New Synonymy in the Haemogamasidae, Laelaptidae and Diplo- gyniidae Indicated by an Examination of Banks' Types of Mesostigma- ta (Acarina). D. E. Johnston 60 Additions to the Bioecology of the New England Tingidae and Piesmidae ( Heteroptera) . N. S. Bailey 63 •Glandular Sources and Specificity of Some Chemical Releasers of Social Behavior in Dolichoderine Ants. E. O. Wilson and M. Papuan 70 Author and Subject Index for Volume 66 79 nei CAMBRIDGE ENTOMOLOGICAL CLUB Officers for 1959-60 President M. Parsons, Harvard University Vice-President A. Bull, Wellesley College Secretary J. J. T. Evans, Harvard University Treasurer F. M. Carpenter, Harvard University Executive Committee E. 0. Wilson, Harvard University N. W. Gillham, Harvard University EDITORIAL BOARD OF PSYCHE F. M. Carpenter (Editor), Professor of Entomology , Harvard U niversity P. J. Darlington, Jr., Head Curator of Recent Insects , Museum of Comparative Zoology W. L. Brown, Jr., Associate Curator of Insects , Museum of Compara- tive Zoology E. 0. Wilson, Associate Professor of Zoology , Harvard University H. Levi, Associate Curator of Arachnology , Museum of Compara- tive Zoology PSYCHE is published quarterly, the issues appearing in March, June, Sep- tember and December. Subscription price, per year, payable in advance: $4.50 to Club members, $5.00 to all other subscribers. Single copies, $1.25. Checks and remittances should be addressed to Treasurer, Cambridge Ento- mological Club, Biological Laboratories, Harvard University, Cambridge, Mass. Orders for back volumes, missing numbers, notices of change of address, etc., should be sent to the Editorial Office of Psyche, Biological Laboratories, Har- vard University, Cambridge, Mass. IMPORTANT NOTICE TO CONTRIBUTORS Manuscripts intended for publication, books intended for review, and other editorial matter, should be addressed to Professor F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. Authors contributing articles over 8 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 $6.00 per page. The actual cost of preparing cuts for all illustrations must be borne by contributors; the expense for full page plates from line drawings is approximately $8.00 each, and for full page half-tones, $10.00 each ; smaller sizes in proportion. AUTHOR’S SEPARATES Reprints of articles may be secured by authors, if they are ordered before, or at the time proofs are received for corrections. The cost of these will be furnished by the Editor on application. The September, 1959 Psyche (Vol. 66, no. 3) was mailed Sep- tember 8, i960. The Lexington Press. Inc., Lexington, Massachusetts William M. Mann 1886-1960 From a photograph taken in October, 1956. Courtesy of the Smith- sonian Institution and Mrs. Lucile Q. Mann. PSYCHE Vol. 66 December, 1959 No. 4 WILLIAM M. MANN “ Each day I would stroll along the beach to a new stream and follow it to a new part of the forest, in a continual state of exultation over the abundant and interesting specimens.” Ant Hill Odyssey The intense enthusiasm with which William Mann undertook the exploration of remote insect faunas was the dominant note in his unusual and distinguished scientific career. As a young man embark- ing on a series of extensive collecting trips to Brazil, Mexico, the West Indies, the Middle East, the South Pacific, and other parts of the world, he openly sought personal adventure as a major reward of scientific endeavor. The “Ant Hill Odyssey”, as he later signified it in the title of his autobiography, was never allowed to stagnate in the physical confinement and routine that sometimes make scholarly enterprise falsely seem less than a great adventure. This spirit he was able to transmit to younger entomologists, and it was responsible for the beginning of the career of more than one young field biologist. There would be little gain in attempting to recall here the events of Mann’s crowded life already told so vividly in his autobiography. It may be noted that most of his entomological field work was con- ducted while he was a graduate student, and then a Sheldon Travel- ling Fellow, at Harvard University during 1911-17. As a student he served as Secretary of the Cambridge Entomological Club and Assistant Editor of Psyche. In 1917 he received a joint appointment in the United States Department of Agriculture and National Mu- seum. In 1925 he fulfilled a life-long dream to become a zoo director when he succeeded Alexander Wetmore as head of the National Zoo- logical Park. His autobiographical account ends with his 1917 ap- pointment, but in fact the entomological odyssev never ended. As director of the national zoo, Mann made several major expeditions abroad to collect living animals and passed up no opportunity to gather insects, especially his beloved ants and ant-guests, on the side. Those privileged to know him in his later years could still sense the full ex- 55 56 Psyche [December citement of this continuing adventure in his wondrous after-dinner anecdotes of field trips around the world. A common story has it that Mann’s faculty sponsor at Harvard, William Morton Wheeler, was at first keenly disappointed when he abandoned a full-time career as entomologist for zoo-keeping but soon became completely recon- ciled by his former student’s obvious genius in the latter role. At the National Zoological Park, Mann was enormously successful. He de- veloped humane, new techniques in zoo culture and was responsible for the introduction of many new animals to zoo life. He was re- nowned for the wit and eloquence with which he sought, and success- fully obtained, the congressional appropriations needed to expand his zoo. His unfailing hospitality was extended to persons from all walks of life, and he had many close friends and an army of warm personal admirers. “Small in stature, puckish, bright-eyed and almost formally unkempt, Dr. Mann possessed a gentle wit which he used sparingly in public — lest it be considered unseemly in a scientist of his ac- knowledged standing. . . He became a familiar figure to thousands of Zoo patrons who brought away with them the charming memory of intimate little chats he was never too busy to hold with the least of his visitors.’’* When he died in his Washington home on October IO, i960, at the age of seventy-four, the Ant Hill Odyssey perhaps seemed to many of his friends no more than a remote chapter in a colorful past. Yet it should not be forgotten that his early work produced major contributions that have actually gained in value with the pass- age of time. Mann’s collections of ants and myrmecophiles, which are the most significant parts of his general collections, are divided chiefly between the Museum of Comparative Zoology and the U. S. National Mu- seum. His entomological publications are based mostly on this ma- terial, a fact that gives them their exceptional value. Mann’s collec- tions, especially those of ants made in the West Indies and South Pacific, were unusually thorough. Few men have been able to write taxonomic papers on tropical insect faunas with such an intimate first- hand knowledge of the ecology and faunal relationships of his study material. As a result, his larger monographs have been little improved on by later work and will undoubtedly remain primary references for years to come. The following bibliography includes as complete a list of Mann’s technical entomological publications as could be assembled at this time. References to his well-known popular article on ants in the National Geographic and to his autobiography are also included. *The Washington Post , October 11, 1960. 1959] Wilson — William M. Mann 57 No attempt has been made to list his articles dealing with zoo culture, travel, and other popular topics. E. O. Wilson Entomological Publications of William M. Mann 191 1 Notes on the guests of some Californian ants. Psyche, 18(1) 127-31. On some northwestern ants and their guests. Psyche, 18(3) :i02- 109. 1912 A third collection of Mallophaga from Alaskan birds. Ent. News, 23:12-17. (With V. L. Kellogg) . Mallophaga from islands off Lower California. Ent. News, 23:56- 65. (With V. L. Kellogg). Parabiosis in Brazilian ants. Psyche, 79 ( 2) 136-4 1. Note on a guest of Eciton hamatum Fabr. Psyche, 29(3) 198- 1 00. List of Histeridae and Buprestidae (Stanford Expedition to Brazil, 1 9 1 1 ) . Psyche, 79(4) :i 18-121. A protective adaptation in a Brazilian membracid. Psyche, 79(5) : 145-147- Literature for 1 91 1 on the behavior of ants and myrmecophiles. J. Animal Behavior, 2(6) : 400-420. 1913 Literature for 1912 on the behavior of ants and myrmecophiles. J. Animal Behavior, 3(6) 1429-445. Mallophaga from Brazilian birds. Psyche, 2o( 1 ) : 1 5-23. (With J. H. Paine). 1914 Some myrmecophilous insects from Mexico. Psyche, 27(5) 1172- 184. The ants of Haiti. Bull. Amer. Mus. Nat. Hist., 3j(i):i-6i. (With W. M. Wheeler). 58 Psyche [December 1915 A new form of a southern ant from Naushon Island, Massachusetts. Psyche, 22 { 2) 151. A cursorial tick. Psyche, 22(2) :6o. Some myrmecophilous insects from Hayti. Psyche, 22(5) :i6i-i66. A gynandromorphous mutillid from Montana. Psyche, 22 ( 5) 1178- 180. 1916 The ants of Brazil (Stanford Expedition to Brazil, 1911). Bull. Mus. Comp. Zool. Harv., 60 { 1 1 ) 1399-490, 7 plates. The ants of the Phillips Expedition to Palestine during 1914. Bull. Mus. Comp. Zool. Harv., £>0(5) 1167-174. (with W. M. Wheeler). 1918 Myrmecophilous insects from Cuba. Psyche, 25 ( 5) 1104- 106. 1919 The ants of the British Solomon Islands. Bull. Mus. Comp. Zool. Harv., 63 ( 7) 1273-391, 2 plates. 1920 A proctotrypid inquiline with Formica exsectoides Forel (Hym.). Proc. Ent. Soc. Wash., 22(3) :59-6o. Ant guests from Fiji and the British Solomon Islands. Ann. Ent. Soc. Amer., 13(1) :6o-6g. Additions to the ant fauna of the West Indies and Central Ameri- ca. Bull. Amer. Mus. Nat. Hist., 42 ( 8) .*403-439. 1921 The ants of the Fiji Islands. Bull. Mus. Comp. Zool. Harv., 64 (5) .*401-499. A new genus of termite guest from Fiji. Psyche, 28(2) 154-56. Three new myrmecophilous beetles. Proc. U. S. Nat. Mus., 59: 547-552. 1922 Notes on a collection of West African myrmecophiles. Bull. Amer. Mus. Nat. Hist., 45: 623-630. Ants from Honduras and Guatemala. Proc. U. S. Nat. Mus., 61 (13) :i-54- 1923 New genera and species of termitophilous Coleoptera from north- ern South America. Zoologica, 3(17) : 323-366. 1959] Wilson — William M. Mann 59 Two new ants from Bolivia. Psyche, 30(1) : 1 3- 1 8. Two serphoid guests of Eciton (Hym.). Proc. Ent. Soc. Wash., 23(9) : 1 8 1- 1 82. A singular habit of sawfly larvae. Psyche, 30(1) :9_I3- (With W. M. Wheeler). 1924 Myrmecophiles from the Western United States and Lower Cali- fornia. Ann. Ent. Soc. Amer., ij( 1) :Sy-g^. Notes on Cuban ants. Psyche, 31 ( 1 ) : 19-23. 1925 Ants collected by the University of Iowa Fiji-New Zealand Ex- pedition. Iowa Studies in Natural History, 1/(4) :5_6. Guests of Eciton hamatum (Fabr.) collected by Professor W. M. Wheeler. Psyche, 32(3) : 1 66- 177. New beetle guests of army ants. J. Wash. Acad. Sci., 15(4) :73~7 7 - 1926 Some new Neotropical ants. Psyche, 33 ( 4-5) 197-107 . Three new termitophilous beetles from British Guiana. Proc. Ent. Soc. Wash., 28 ( 7) : 1 5 1-1 55. New Neotropical myrmecophiles. J. Wash. Acad. Sci., 16(16): 448-455. 1928 A new Microdon from Panama. Psyche, ^5(3) : 168- 170, 1929 Notes on Cuban ants of the genus Macromischa ( Hymenoptera : Formicidae). Proc. Ent. Soc. Wash., j/(8) : 1 6 1 - 1 66. i93i A new ant from Porto Rico. J. Wash. Acad. Sci., 2/(17) ^40-441 . 1934 Stalking ants, savage and civilized. National Geographic Mag., 66 2) 1171-192. 1935 Two new ants collected in quarantine. Psyche, 42(1) .*35-37. 1948 Ant Hill Odyssey. Little, Brown. (Autobiography). SOME NEW SYNONYMY IN THE HAEMOGAMASIDAE, LAELAPTIDAE AND DIPLOGYNIIDAE INDICATED BY AN EXAMINATION OF BANKS’ TYPES OF MESOSTIGMATA (ACARINA) By Donald E. Johnston Institute of Acarology, Department of Zoology, University of Mainland During the course of a study of certain of the Banks mite types in the collection of the Museum of Comparative Zoology at Harvard College some cases of obvious synonymy were found. Two of these cases sink Banks’ names; the remainder result in the establishment of his names as senior synonyms. In addition to those of Banks, the types of Ewing’s, Furman’s and Hicks’ species (all in U.S. National Museum) were examined in connection with this work. I am grateful to Drs. H. W. Levi and W. L. Brown for arranging for me to study the Harvard collection and for their many kindnesses during my visit. The following lectotvpe designations are not as precise as one would wish. T he Banks types in the Museum of Comparative Zoology are not numbered and most names are represented by cotypes. All of the suspected and certain type slides are arranged alphabetically according to species and kept as a unit in the Arachnida collection of the Museum. Thus the lectotypes designated herein will be found labeled as such in their proper place in the alphabetical filing scheme. It may also be noted here that some of Banks’ species (none of those treated herein) are represented by cotypes in both the MCZ and the Acarina collection of the U.S. National Museum. The same is true of Jacot’s material which has been divided evenly between these museums. Haemogamasidae Haemogamasus longitarsus (Banks) Comb. Nov. Lae laps longitarsus Banks, 1910. Proc. Ent. Soc. Wash. 12 (1) 15. Ex mole’s nest, Falls Church, Virginia. Lectotype in MCZ. Haemogamasus barberi Ewing, 1925* Proc. Biol. Soc. Wash. 38 : 140. Ex nest of small mammal; Maryland shore of Potomac River, near Plummer Island. Type in U.S.N.M. new synonymy. Haemogamasus microti Ewing, 1925* Proc. Biol. Soc. Wash. 38: 141. Ex Microtus pennsylvanicus , Bronxville, New York. Type in U.S.N.M. [ — barberi according to Keegan, 1951:249] Remarks: The type series of Laelaps longitarsus consisted of eight 60 1959] Johnston — Haemogamasidae 61 specimens. These have been remounted individually and a female is here designated as lectotype (indicated on slide label). E nine l a ps stab ill arts ( Koch ) Gamasus stabularis Koch, 1836. Crust, Myr. Arachnid. Deutsch- lands, Fasc. 27, No. 1 Host unknown; Regensburg. Type unknown. Laelaps pedalis Banks, 1909. Proc. Ent. Soc. Wash. 11 (3) :i36. Ex chipmunk; Guelph, Ontario, Canada. Type in MCZ. new SYNONYMY. Laelaps prophetic us Banks, 1909. Ibid 1136 Ex groundhog; Guelph, Ontario. Lectotype in MCZ. new synonymy. Remarks: The type slide of L. propheticus contained one female E. stabularis (here designated lectotype) and two Haemolaelaps glasgowi (Ewing). Laelaps pedalis is represented by a single female. The concept of E. stabularis with which Banks’ names are synonymized is that of Strandtmann and Wharton (1958:127) and Bregatova (1956:100). Laelaptidae Androlaelaps frontalis (Banks) Comb. Nov. Gamasus frontalis Banks, 1910. Proc. Ent. Soc. Wash. 12 (3) 13. Ex Perognathus ; Los Palmos, 3500 feet, Santa Rosa Mountains, California. Lectotype in MCZ. Androlaelaps sinuosa Furman, 1954. Pan-Pacific Ent. jo (2) :i20. Ex Pergognathus sp., probably fallax fallax , at Pigeon Pass, River- side County, California. Type in U.S.N.M. new synonymy. Remarks: The type slide bore four females and two males of this striking laelaptid. These have been remounted individually and a female is here designated lectotype (indicated on slide label). Banks’ placement of this species in Gamasus (presumably because of the large size and calcarate femora of legs II) is undoubtedly the reason for its having been overlooked by workers in the Laelaptidae. Diplogyniidae Neolobogynium americana (Banks) Comb. Nov. Celaenopsis americana Banks, 1906. Proc. Ent. Soc. Wash. 7: 137. Ex Histerid beetle (Elololepta sp.) at Indianapolis, Indiana. Lectotype in MCZ. Neolobogynium lateriseta Hicks, 1957* Proc. Iowa Acad. Sci. 64: 616. Host unknown; Ames, Iowa. Type in U.S.N.M. new syn- onymy. 62 Psyche [December Remarks: Banks’ type slide contained 18 adults of both sexes of this common mite associate of Uololepta ( Histeridae) . These have been remounted and a female is here designated lectotype (indicated on slide label). References Cited Bregatova, N. G. 1956. Gamasid Mites (Gamasoidea) . Acad. Sci. U.S.S.R., Moskow. 247 pp. [in Russian.] Keegan, H. L. 1951. The mites of the subfamily Haemogamasinae (Acari: Laelaptidae) . Proc. U. S. Nat. Mus. 101: 203-268. Strandtmann, R. W. and G. W. Wharton. 1958. A manual of mesostigma- tid mites parasitic on vertebrates. Contribution 4, Institute of Acarology, University of Maryland, i-xi-j- 1-330 + pis. 1-69. ADDITIONS TO THE BIOECOLOGY OF THE NEW ENGLAND TINGIDAE AND PIESMIDAE (HETEROPTERA)1 By Norman S. Bailey Bradford Junior College, Bradford, Mass. Intermittent summer collecting during the past ten years has added to my collection a considerable number of noteworthy records. These include additional stations for species previously reported from only a few localities within a particular state and for some of the less well- known species in New England, new records for the occurrence of six species previously un reported from one or more of the New Eng- land states, and a few new host plant records. This is the first of a series of papers now in progress which will supplement my studies of the Tingidae of New England ( 1951 ). For ease of reference, therefore, it follows the pattern of that study in the sequence of genera and of species. (Bailey, 1951, page 5). All collections were made by the author unless otherwise specified. Family Piesmidae Piesma cinerea (Say) Shelton, Connecticut, August 2, 1951; Aina, Maine, August 15, 1954 and August 28, 1956. Found on a weedy Amaranthus (probably retroflexus L. ) on each occasion. In 1917 Professor Parshley reported this species from ocean drift at Beach Bluff, Massachusetts, and stated, “Not found heretofore north of Rhode Island”. Since then it has become well-known from this state. The records from east-central Maine extend its range ap- preciably. Aina is a small town adjoining Wiscasset in the Sheepscot River Valley about fifty miles east-northeast of Portland. Since published records for Connecticut are few, note is made of the Shelton collection. Family Tingidae Leptoypha rnutica (Say) Windham, New Hampshire, August 11, 1958. A large collection of adults, many teneral, a few nymphs, and some exuviae was made from suppressed ash ( Frcixinus) and ash suckers Acknowledgement is hereby gratefully made of a Sigma Xi • — RESA Re- search Fund grant and Grant No, G5477 from the National Science Founda- tion, which make these studies possible. 63 64 Psyche [December growing on a dry, rocky hillside in a stand of young mixed hard- woods. This is, I believe, the third New Hampshire record for this small lace bug. Galeatus peckhami (Ashmead) Katahdin Trail, Maine, August 14, 1959; East Madison, New Hampshire, July 25, 1958. Extended search was finally rewarded by these two collections from roadside asters ( Aster macrophyllus L.) growing on dry, sunny banks. Though this host plant is common in northern New England, frequent search has provided only these two records. Corythucha marmorata (Uhler) is more commonly found on this aster. Gargaphia til'iae (Walsh) New Milford, Connecticut, August 2, 1951 ; Aina, Maine, August 29, 1956; Carroll, Maine, August 14, 1959; Petersham, Massachu- setts, July 17, and 18, 1951; Conway, New Hampshire, July 25, 1958; Eaton, New Hampshire, July 25, 1958. In general this arboreal species may be found on its host plant Tilia. Since few records have been published, those noted will extend our knowledge of its distribution. All the collections were made from Tilia. Corythaica bellula Bueno Aina, Maine, August 2, 1953, August 28, 1956, and September 9, 1958; Cedar Grove, Maine, July 11, 1953; Petersham (Harvard Forest), Massachusetts, July 16 and 17, 1951; Conway, New Hampshire, July 25, 1958 (sweeping) ; Eaton, New Hampshire, July 25, 1958. Each collection was made by hand from Panicum, except in the instance noted. The Aina station was a wood road through a mature stand of Pinus strobus L. where the shade was almost constant but not intense and at a point where the road was beginning to rise at the base of a fairly sharp slope. This species was also' seen, but not col- lected, on July 11, 1953, on Panicum growing on a little used, open drive within a stone’s throw of the shore at Medomak, Maine. The Cedar Grove collection was from Panicum growing on a dry bluff overlooking the Kennebec River. These are the first Maine records. In Bradford, Massachusetts, six collections of C. bellula Bueno were made from Panicum by hand and two by sweeping in 1957. Most of these were taken from an area of less than four square yards under a small red maple ( Acer rubrum L.) where the Panicum was prominent and the total vegetation too sparse to cover the hard-packed 1959] Bailey — Tingidae and Piesmidae 65 soil. These eight collections yielded 14 females, 4 males, and 8 nymphs. In 1958, six collections in the same area gave 6 females, 10 males, and 14 nymphs. The first specimens were taken on June 9th in 1957 and the last for the season on September 7th in 1958. Nymphs were taken as early as June 12th and as late as August 31st. Teneral adults occurred from as early as June 22nd until as late as August 17th. These data confirm the opinion that C. bellula Bueno should be found throughout New England in areas ecologically suitable. Stephanitis globulifera (Matsumura) West Hartford, Connecticut, August 25, 1958. They were abun- dant on Pieris japonica (Thunberg) Don used as foundation plants in front of a fairly new house in a recent development. This species first reported from North America in 1950 (Bailey) is apparently well-established in Connecticut, but I have yet to collect it elsewhere. Stephanitis pyrioides (Scott) West Newbury (Cherry Hill), Massachusetts, August 20, 1958. Nymphs and adults were found on young stock of several deciduous azalea species and hybrids growing in a lath house. Stephanitis rhododendri Horvath Petersham (Harvard Forest), Massachusetts, July 17 and 18, !95!. The first collection was made from Kahnia latifolia L., a frequent host. The next day a few were found on K. angustifolia L. Al- though this is a very common shrub in eastern Massachusetts and Maine and, although it is on record as a host for this lace bug, this is my only collection from lambkill. Corythucha bellula Gibson Katahdin Trail, Maine, August 14, 1959; Gorham, New Hamp- shire, July 21, 1958. This is the first record of the species from New Hampshire where it was found on roadside Cory bus. Note below that C. coryli O. and D. also occurs in this state. The ecology of these two species in their contact zone should be a fruitful study since both utilize the same host. The Maine collection noted is a large one of particular interest since they were all taken from Salix. Corythucha caryae Bailey Scituate, Rhode Island, September 23, 1958. This interesting range extension of a species previously collected only from a relatively limited area in eastern New Hampshire and 66 Psyche [December Massachusetts was the outstanding result of a one-day collecting trip into Rhode Island. They were found on Carya on the edge of a wooded area beside Route 14 and near the reservoir. Corythucha coryli O. and D. Monroe, Connecticut, August 2, 1951; Shelton, Connecticut, August 27, 1958; Epping, New Hampshire, September 20, 1958; Diamond Hill State Park, Rhode Island, September 23, 1958. My collections now extend the range of this species into three more of the New England states for which there have been no published records. They were all taken from Corylus growing along the road- side. Corythucha cydoniae (Fitch) Newcastle, Maine, August 7, 1953; Haverhill (Ward Hill), Mas- sachusetts, July 27, 1957 and August 23, i960; Diamond Hill State Park, Rhode Island, September, 23, 1958. This species can now be added to the faunal lists of two more New England states. Since it infests shrubs and small trees of several rosaceous genera that are widely distributed in the Northeast, the single reported occurrence north of Massachusetts suggested a wider distribution and also raised the question of limiting ecological factors. The wider range is now established but the ecological conditions con- trolling it surely deserve investigation. I have collected rather widely and, in a few areas, intensively in Maine with only the record cited above for this species. There it was feeding on native Crataegus in large numbers. The single Rhode Island record, on the contrary, probably indi- cates lack of local interest in Tingids. The Massachusetts records are given to report another host for C. cydoniae. Cotoneaster hupehensis Rehder and Wilson, used in a permanent shrub border in a local nursery was found to be heavily infested. About twenty yards away is a large clump of tall Amelan- chier , also a part of the landscape arrangement, similarly infested. The lace bugs were breeding on both hosts. The eighth edition of Gray's Manual of Botany (Fernald, 1950) considers Pyracantha coccinea Roemer (a previously reported host) a synonym for Cotoneaster pyracantha (L.) Spach while the 1949 revised edition of the Manual of Cultivated Plants (L.H. Bailey) maintains the distinctness of the genus Pyracantha. This leaves me with a question for the plant taxonomists to decide. If not another genus, we at least note another rosaceous species as a host for C. 1959] Bailey — Tingidae and Piesmidae 67 cydoniae which thrives on many native and cultivated forms in this plant family. Corythucha marmorata (Uhler) Bradford, Massachusetts, July 6, 1957; Haverhill (Ward Hill), Massachusetts, July 27, 1957. This species ranges throughout North America. In New England it feeds exclusively on a diversity of herbaceous composites. Apparent- ly one of America’s most successful Tingids in terms both of its wide distribution and of its local abundance, it is the only species of the fifteen in the genus occurring in New England that is neither arboreal nor thamnophilous. Its catholic preference for ubiquitous composites undoubtedly accounts in no small measure for its success. Consequently, it was not surprising to find it thriving on two species previously unlisted. The Bradford observation was made on Rudbeckia serotina Nuttall, the common Black-eyed Susan or Yellow Daisy. On Ward Hill they were found on the cultivated Globe Thistle, Echinops sphaerocephalus L. Corythucha mollicula O. and D. Monroe, Connecticut, August 2, 1951; Newtown, Connecticut, August 2, 1951; Aurora, Maine (collected by Dr. A. G. Humes), July 13, 1950; Rockport, Maine, September 10, 1958; Sheepscot, Maine, August 6, 1953; Petersham (Harvard Forest), Massachu- setts, July 18, 1951; Diamond Hill State Park, Rhode Island, Sep- tember 23, 1958. Probably again because of lack of collector interest, few records have been published for this species. Therefore, these, all from Salix , are cited. Corythucha pruni O. and D. Brookfield, Connecticut, August 2, 1951; Monroe, Connecticut, August 2, 1951; Shelton, Connecticut, August 27, 1958; Mattatuck, Connecticut, August 28, 1958; Aina, Maine, August 28, 1956 and September 9, 1958; Cedar Grove, Maine, September 15, 1957 and May 28, 1959; Dresden, Maine, June 22, 1957; Mexico, Maine (collected by Mr. J. J. B. Anderson), September 7, 1959; New- castle, Maine, August 7, 1953; South Bristol, Maine, August 24, 1956; South Cushing, Maine, September 10, 1958; Epping, New Hampshire, September 20, 1958; Diamond Hill State Park, Rhode Island, September 23, 1958; Waterman, Rhode Island, September 23, 1958. 68 Psyche [December The Maine records cited establish the frequent occurrence of this species in the central coastal region and in other parts of that state. Another locality is added to the few published New Hampshire rec- ords. Rhode Island and Connecticut stations are given for the first time. Of particular interest is the fact that the specimens from the two southernmost New England states suggest that typical Corythucha pruni O. and D. and typical C. associata O. and D. may represent extreme forms which have differentiated in the recent past from a common population. I he latter has not yet been collected in Mas- sachusetts nor further north. Until now, C. pruni had not been re- ported from southern New England. Both species restrict their feed- ing entirely to Prumis serotina Ehrhart. The collections noted above were all from this host. Two collections made in Connecticut in I951 aroused my curiosity. In both, specimens of the two species were found on the same host plants. On recent critical examination of this material, I found that the Brookfield collections consisted of 26 specimens. Three of these were, on basis of hood size and form, definitely C. pruni (2 males and a female), 22 were C. associata, and one was questionably C. associata because the top of the hood was missing. Among the associata forms there was some variation in the height and width of the hood. The Monroe collection made the same day consisted of 38 that were surely C. pruni and a single female of C. associata. More recently a collection from Mattatuck produced 9 males of C. pruni and one female of C. associata. In Shelton one C. pruni male was found on a Prun.us and another host plant in that town yielded 100 adults of C. associata. Close study of these two species in Connecticut should answer some important questions. Except for the marked differences in their hood forms and, more significantly, the claspers of the males, they are very similar. In size and coloration they are almost identical. Frequent collections in several localities in this contact zone throughout one or two seasons would give valuable population information and speci- mens for careful comparison of structural details. As previously mentioned, a similar situation involving the species C. bellula Gibson and C. coryli O. and D. whose primary host is Cory lus may exist from central Maine and New Hampshire west- ward. Again we find two species of similar size with minor mor- phological distinctions. Similarly intriguing is the relationship be- tween our two New England species whose host is commonly Alnus. Here, however, there are greater differences in size in addition to the 1959] Bailey — Tingidae and Piesmidae 69 differences in structural details between the two and, consequently, their kinship is possibly more remote. Corythucha ulrni O. and D. Brookfield, Connecticut, August 2, 1951. Published Connecticut records are few. In my experience, this species is only rarely found even where elms are common. Literature Cited Bailey, L. H., 1949. Manual of Cultivated Plants. Macmillan, New York. Bailey, Norman S., 1950. An Asiatic Tingid New to North America. Psyche, 57:143-145. 1951. The Tingoidea of New England and Their Biol- ogy. Entomologica Americana, 31 (n.s.) :1-14Q. Fernald, M. L., 1950. Gray’s Manual of Botany (Eighth Edition). Ameri- can Book Company, Boston. Parshley, H. M., 1917 a. Insects in Ocean Drift. Canadian Entomologist, 49 (2) :45-48. 1917 b. Fauna of New England, 14. List of the Hemip- tera-Heteroptera. Boston Society of Natural History, Occasional Papers, 7:1-125. GLANDULAR SOURCES AND SPECIFICITY OF SOME CHEMICAL RELEASERS OF SOCIAL BEHAVIOR IN DOLICHODERINE ANTS1 By Edward O. Wilson2 and Mario Pavan3 Introduction Species of the ant subfamily Dolichoderinae are excellent subjects for the experimental analysis of chemical communication. The group is advanced evolutionarily, and large, complex societies are the rule. During foraging, workers of many species form long, conspicuous columns tightly bound to persistent odor trails. When disturbed they void volatile secretions that seem likely to function, at least in part, as olfactory releasers of alarm behavior. Furthermore, the dolicho- derines have recently been the object of extensive biochemical research by Pavan, Cavill and their associates (cf. Pavan, 1950; Pavan and Ronchetti, 1955; Pavan and Trave, 1958; Cavill, Ford, and Locks- ley, 1956; Cavill and Locksley, 1957). Several terpenoids including the previously unknown iridomyrmecin, iso-iridomyrmecin, and irido- dial, have been identified as components of the anal gland secretion of various species. T he availability of these substances in purified form has made possible for the first time the precise behavioral assay ol natural secretory products in ants. Trail Substances Applying the artificial trail technique used earlier by one of us (Wilson, 1959) in studies of the myrmicine Solenopsis saevissima , an attempt was made to determine the glandular source of the trail sub- stance in workers of Iridomyrmex humilis Mayr. Groups of workers from laboratory colonies (grown from stocks originating from Baton Rouge, Louisiana) were attracted to drops of 1 M sucrose solution, about 6 mm in diameter, on glass feeding tables. Selected body parts of freshly killed workers were made into artificial trails directed at the feeding workers. The number of workers following the trails be- yond the trail midpoint, i.e., 15 cm. from the sucrose bait, were re- corded. When positive responses were obtained, the duration of biological activity was timed to the nearest half-minute. Preliminary experiments showed that the trail substance is limited to the abdomen, ^ased on research supported in part by a grant from the U. S. National Science Foundation. ^Biological Laboratories, Harvard University, Cambridge, Mass., U. S. A. 3Istituto di Anatomia Comparata and Istituto di Entomologia Agraria delPUniversita di Pavia, Italy. 70 1959] Wilson and Pavan — Chemical Releasers in Ants 71 smears from other body parts proving consistently inactive. This re- sult was not unexpected, since humilis workers can be clearly seen to drag the posterior portion of the abdomen over the ground while in the act of laying trails. Five organs in the abdomen are capable of emptying glandular se- cretions to the outside: (1) the hind-gut; (2) the paired “true” poison glands, with a large reservoir, the poison vesicle; (3) Dufour’s gland (also called the accessory gland) ; (4) the anal glands, opening just above the anus; (5) the glandular “ventral organ” recently dis- covered by Pavan ( 1955). In a series of experiments, the five organs were dissected out of freshly killed workers, washed in insect Ringer’s solution, and bioassayed with artificial trail tests. The results, sum- marized in table 1, show that the bulk, and perhaps the entirety, of fhe trail substance resides in the ventral organ. The occasional weak Table 1. Results of artificial trail assay of selected abdominal organs of Iridomyrmcx humilis workers. Organ Number of tests Number of positive responses* Number of workers responding : range (with mean) Duration of re- sponses in minutes: range (with mean) hind gut 6 t) 0-2 (M< 1 ) 0-1/2 (M< 1/2) true poison glands, plus reservoir 6 0 0-2 (M<1) 0-1/2 (M< 1/2) Dufour’s gland 8 2 0-30 (M = 8) 0-4 (M = l) anal gland S 1 0-15 (M = 6) 0-1 ( M — 1 ) ventral organ 7 ? 65-339 (M=134) 2-9 (M — 5) * Positive responses are defined, arbitrarily, as the crossing of the trail mid- point by 10 or more workers. responses to the contents of the Dufour’s and anal glands may not be biologically significant, i.e., may not indicate the presence of sub- stances normally secreted into the trail, since responses of comparable magnitude have been obtained with pure methylheptenone, a sub- stance not known to occur in humilis workers. In a second series of experiments, the ventral organ and anal glands of Iridomyrmcx pruinosus (Roger)4 were assayed. As in humilis , the ventral organ smears caused strong trail following while re- Tollected at Baton Rouge, Louisiana. 72 Psyche [December spouses to anal glands were negligible. Furthermore, artificial trails made with 2-heptenone, the anal gland secretion of pruinosus (M. S. Blum, in litt. ) , caused negligible responses. In a third series of experiments, the hind-gut, poison apparatus, Dufour’s gland, and ventral organ were assayed in a laboratory colony of the primitive dolichoderine Monads bispinosa (Olivier).* The unusually restless and aggressive nature of the Monads workers required the following modification in procedure. Straight trails 75 cm. in length were drawn down the middle of a clean glass plate. Three workers were then released, one at a time, and allowed to wander over the glass plate, crossing and re-crossing the trail. A positive response was recorded when the worker persistently followed the trail to its end. Of 18 workers thus exposed to trails made suc- cessively from 6 ventral organs, 17 gave positive responses. The re- sponses of workers in triplicated control experiments using the other abdominal organs were all negative. After the chief source of the odor trails in the three dolichoderine species had been located, bioassays were made across species, in order to determine the extent of species-specificity of the trail substances. As shown in table 2, each species appears to have a different substance. Table 2. Intra- and interspecific trail-substance tests employed in the present study. Only intraspecific tests produced positive responses ( + ). The results of all interspecific tests employed were negative ( ). Interspecific tests not attempted are Source Monads indicated by a question mark. Recipient Liometopum lridomyrmex lridomyrmex T apinoma bispinosa occidental humilis pruinosus sessile M. bispinosa + ? — ? L. occidentale ? + ? — /. humilis - + — — 1. pruinosus ? ? — _L 1 — T. sessile j> ? + Further, the Dufour’s glands and ventral organs of Monads bispinosa were assayed with laboratory colonies of the myrmicine ants Cremato- gaster lineolata (Sav), So/enopsis geminata (Fabr.), and S. saevissi- nia (Fr. Smith). The ventral organ induced no detectable response in these species. The Dufour’s gland caused no response in the Crematogaster but, quite unexpectedly, caused strong trail-following in Solenopsis saevissima! In fact, single Monads glands consistently drew out approximately the same number of S. saevissima workers as single saevissima glands. It was also determined that a single Monads Collected at Palmar, southern Costa Rica. 1959] Wilson and Pavan — Chemical Releasers in Ants 73 Dufour’s gland caused a weaker response in S. getnmata , well below the intensity caused by a geminata gland and approximately the same as that caused by a saevissima gland (see Wilson, i960). These results are interpreted as indicating the chemical identity, or near- identity, of the Monacis bispinosa and Solenopsis saievissima Dufour’s secretions. Curiously, the Solenopsis use the secretion as a trail sub- stance, but the Monacis do not. The function of the secretion in Monacis remains unknown. Some further comment is needed concerning the organ we are here referring to as the ventral organ. The gland was discovered in Iridomyrmex humilis by Pavan ( 1955) and described in greater detail by Pavan and Ronchetti (1955). It is a paddle-shaped organ resting on the ventral body wall just over abdominal sternite VI (sternite IV of gaster) . It opens posteriorly via a neck-like constriction between the VI and VII abdominal sternites. The opening is fitted with an unusual beak-shaped deformation of the posterior border of sternite VI, a structure seemingly designed to guide the effluence of the gland contents. Until the present experimental work, the function of the gland remained unknown, although Pavan and his associates suggested that it produced either the trail substance or a secondary defensive secretion. Additional morphological studies by Miradoli Zatti and Pavan (1957) disclosed the presence of a closely similar gland in the primitive dolichoderine genera Aneuretus and Leptomyrmex. In the course of subsequent work we have noted its presence in Monacis bispinosa and Hypoclinea doriae Emery as well. The ventral organ appears to be peculiar to the Dolichoderinae and Aneuretinae and hence can serve as a valuable diagnostic character for these two closely allied groups. If it serves as the source of the trail substance in all of the dolichoderine tribes (and in the ancestral aneuretines) , as the preliminary evidence suggests, it can be interpreted as a remarkable example of a “social organ” evolved de novo with the specific func- tion of mediating worker communication. Alarm Substances When colonies of Tapinoma sessile , Liometopu?n occidental , and Monacis bispinosa are disturbed by opening the nest, workers release considerable quantities of secretions from their anal glands which can be easily smelled by the human observer. When the anal glands are removed from freshly killed workers of these species and their con- tents released in the presence of groups of resting live workers, the latter are thrown instantly into a typical alarm frenzy, indistinguish- 74 Psyche [December able from that caused by direct mechanical disturbance. Other body parts tested in this manner do not induce the same reaction. Al- though the volatile components of the anal glands of each species pro- duce a distinctly different smell to humans, there is evidently no species-specificity shown in the alarm response. Unlike the complete specificity shown in responses to trail substances, squashes of anal glands of each species produce approximately the same intensity of alarm curiosity in workers of the other two species as in its own. These results can be interpreted as indicating that either a common component exists in each of the anal secretions or else, as seems more likely, the secretions differ but the reactions are not species-specific. In contrast, disturbed workers of Iridomyrmex humilis do not re- lease quantities of volatile substances sufficient to be detected by the human observer. Moreover, they do not respond with more than mild excitement to single anal-gland squashes from sister workers or the other three dolichoderine species. Pavan and Trave (1958) have shown that the volatile components of anal gland secretion of Tapinoma nigerrimum (Nylander) are methylheptenone (C8H140) and propyl-isobutyl-ketone (C8HieO). Since our initial experiments suggested that the action of alarm sub- stances is not species-specific, it was not surprising to find that small quantities of the two nigerrimum substances, manufactured synthet- ically and presented in separate tests, caused intense alarm behavior in Tapinoma sessile. No attempt has been made to map the full range of compounds that will induce the alarm frenzy in the dolichoderines that communicate alarm by anal secretions. However, the following preliminary study will show that not all volatile ant secretions have this effect. A simple olfactometer test designed to measure mass re- sponse in laboratory colonies of Solenopsis saevissima (cf. Wilson, i960) was applied to a small (300-worker) colony of Tapinoma sessile. The tested compound was allowed to evaporate from a 15 mm2 film into a nest inlet into which air was being gently drawn. The maximum increase, if any, of workers outside the nest was then measured during the ensuing five minutes. Each substance was tested three times. Tests were spaced at least two hours apart to reduce habituation, and substances were presented in irregular order to pre- vent special sequential effects. Formic acid, a common defensive abdominal secretion of formicine ants, caused either no visible re- sponse or very mild excitement in the nest, and increased the outside worker force by only 3-8. Concentrated Dufour’s gland secretion of Solenopsis saevissima , which induced a mass exodus of saevissima 1959] Wilson and Pavan — Chemical Releasers in Ants 75 workers in similiar tests, caused in Tapinoma sessile either no visible effect or mild excitement in the nest, and increased the outside worker force from minus- 1 to 9. A mixture of nepetalactone (oil of catnip) and oil of peppermint caused mild excitement in the nest and an exodus of 2 to 24 workers. Propyl-isobutyl-ketone caused mild to in- tense excitement in the nest and an exodus of 12 to 2 3 workers; many of the workers were in maximum intensity of excitement and ranged widely over the foraging table. Methylheptenone caused mild to intense excitement in the nest, and an exodus of 4 to 69 workers, many of which were maximally excited and ranged widely over the foraging table. In one trial using methylpeptonone, a queen, several callows, and other workers carrying brood also left the nest tempo- rarily. In two subsequent experiments, small Tapinoma sessile colonies were exposed in the manner just described to prolonged doses of methytheplenone. In both cases, excitement within the nest and individual worker exodus declined to about the normal level within 10 minutes. Shortly afterward, however, workers began mass move- ments along odor trails to secondary shelters away from the main nest. Within 90 minutes almost the entire colony had emigrated to the new nest sites, leaving behind only a few workers in the main nest. Summary In Iridornyrmex humilis Mayr, I. pruinosus (Roger), and Monacis hispinosa (Olivier), representing two genera phylogenetically far apart in the Dolichoderinae, the principal or exclusive source of the odor trail is the ventral organ, an exocrine gland opening behind abdominal sternite VI. Cross-species tests were made with the odor trails of these species and of Liometopum occidentale Emery and Tapi- noma sessile Say. The trail-substances were found to be species- specific. The ventral organ is peculiar to the Dolichoderinae and the ancestral Aneuretinae, and it is interpreted as an example of a “social organ’’ that has been evolved de novo as part of the colonial communi- cation system. Volatile alarm substances are produced in the anal glands of each of the species except Iridornyrmex humilis . These releasers are not species-specific. The volatile anal secretions of Tapinoma nigerrimum , methylheptenone and propyl-isobutyl-ketone, when tested with work- ers of Tapinoma sessile, produced typical alarm behavior. Prolonged exposure induced colony emigration, which was then organized by the laying of odor trails. 76 Psyche [December References Cavill, G. W. K., D. L. Ford, and H. D. Locksley. (1956). The chemistry of ants. I. Terpenoid constituents of some Australian Iridomyrmex species. Australian J. Chem., 9 (2) : 288-293. Cavill, G. W. K. and H. D. Locksley. (1957). The chemistry of ants. II. Structure and configuration of iridolactone (isoiridomyrmecin) . Australian J. Chem., 10 (3) : 352-358. Miradoli Zatti, M. A. and M. Pavan. (1957). Studi sui Formicidae. III. Nuovi reperti dell’organo ventrale nei Dolichoderinae. Boll. Soc. Ent. Ital., 87 (5-6) : 82-87. Pavan, M. (1950). Potere Insetticida della iridomyrmecina e significato della sostanza vella biologia di Iridomyrmex humilis Mayr. La Ricerca Scientifica, 20 (12): 1853-1855. (1955). Studi sui Formicidae. I. Contribute alia conoscenza degli organi gastrali dei Dolichoderinae. Natura, 46: 135-145. Pavan, M. and G. Ronchetti. (1955). Studi sulla morfologia esterna e anat- omia interna dell'operaia di Iridomyrmex humilis Mayr e richerche chi- miche e biologiche sulla iridomirmecina. Att. Soc. Ital. Sci. Nat., 94 (3-4) : 379-477. Pavan, M. and R. Trave. (1958). Etudes sur les Formicidae. IV. Sur le venin du Dolichoderide T dpinoma nig errimum Nyl. Insectes Sociaux, 5 (3): 299-308. Wilson, E. O. (1959). Source and possible nature of the odor trail of fire ants. Science, 129: 643-644. ( 1960). Chemical communication among workers of the fire ant Solenopsis saevissima (Fr. Smith). Animal Behaviour (in press). CAMBRIDGE ENTOMOLOGICAL CLUB A regular meeting of the Club is held on the second Tuesday of each month October through May at 8:00 p. m. in Room B-455, Biological Laboratories, Divinity Ave., Cambridge. Entomologists visiting Boston are cordially invited to attend. BACK VOLUMES OF PSYCHE The Cambridge Entomological Club is able to offer for sale the following volumes of Psyche. Volumes 3, 4, 5, 6, 7, 8, each covering a period of three years. $8.00 each. Volumes 10, 14, 17 to 26, each covering a single year, $2.00 each. Volumes 27 to 53, each covering a single year, $2.50. Volumes 54 to 65, each covering a single year, $3.00. Volume 66, covering a single year, $5.00. Some other volumes, lacking certain issues, are also available (information upon request). Orders for 2 or more volumes subject to a discount of 10%. Orders for 10 or more volumes subject to a discount of 20%. All orders should be addressed to F. M. Carpenter, Editor of Psyche, Biological Laboratories, Harvard University, Cambridge, Mass. 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 38, Mass. PSYCHE A Journal of Entomology Volume 67 1960 P. J. Darlington, Jr. H. Levi Editorial Board Frank M. Carpenter, Editor W. L. Brown, Jr. E. O. Wilson Published Quarterly by the Cambridge Entomological Club Editorial Office: Biological Laboratories 16 Divinity Ave. Cambridge, Mass., U. S. A. The numbers of Psyche issued during the past year were mailed on the following dates: Vol. 66, no. 4, Dec., 1959: December 28, 1960 Vol. 67, no. 1-2, March-June, 1960: February 16, 1961 Vol. 67, no. 3, Sept., 1960: April 13, 1961 PSYCHE A JOURNAL OF ENTOMOLOGY Established in 1874 Vol. 67 March-June, i960 Nos. 1-2 CONTENTS The Genus Pseudosinella (Collemboia : Entomobryidae) in Caves of the United States. K. Christiansen 1 The Immature Stages of Boriomyia fidelis (Banks) with Taxonomic Notes on the Affinities of the Genus Boriomyia (Neuroptera: Hemero- biidae). E. G. MacLeod 26 The Female of Inocellia pilicornis Carp. (Neuroptera: Inocelliidae) F. M. Carpenter 41 A Note on the Prey and a Nesting Site of Cerceris truncata Cameron. ( Hymenoptera : Sphecidae: Philanthinae ) . F. G. fVerner 43 The Type Species of the Ant Genus Enrhopalthrix. IV. L. Brown, Jr. and JV. IV. Kern pf 44 CAMBRIDGE ENTOMOLOGICAL CLUB Officers for 1959-60 President M. Parsons, Harvard University Vice-President A. Bull, Wellesley College Secretary J. J. T. Evans, Harvard University Treasurer F. M. Carpenter, Harvard University Executive Committee E. 0. Wilson, Harvard University N. W. Gillham, Harvard University EDITORIAL BOARD OF PSYCHE F. M. Carpenter (Editor), Professor of Entomology , Harvard University P. J. Darlington, Jr., Head Curator of Insects, Museum of Com- parative Zoology W. L. Brown, Jr., Associate Curator of Insects , Museum of Compara- tive Zoology E. 0. Wilson, Associate Professor of Zoology , Harvard University H. Levi, Associate Curator of Arachnology, Museum of Compara- tive Zoology 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, $5.00 to all other subscribers. Single copies, $1.25. Checks and remittances should be addressed to Treasurer, Cambridge Ento- mological Club, Biological Laboratories, Harvard University, Cambridge, Mass. Orders for back volumes, missing numbers, notices of change of address, etc., should be sent to the Editorial Office of Psyche, Biological Laboratories, Har- vard University, Cambridge, Mass. IMPORTANT NOTICE TO CONTRIBUTORS Manuscripts intended for publication should be addressed to Professor F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. Authors contributing articles over 8 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 $6.00 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, $18.00 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 1959 Psyche (Vol. 66, no. 4) was mailed December 28, i960. The present issue of Psyche (Volume 67, nos. 1-2) is published with the aid of a National Science Foundation Grant (G 15817) to the Cambridge Entomological Club. The Lexington Press, Inc., Lexington, Massachusetts PSYCHE INDEX TO VOL. 67, i960 INDEX TO AUTHORS Broken, W. L., Jr. and W. W. Kempf. The Type Species of the Ant Genus Eurhopalthrix. 44 Carpenter, F. M. A Triassic Odonate from Argentina. 71 Studies on North American Carboniferous Insects. I. The Protodonata. 98 The Female of Inocellia pilicornis Carp. (Neuroptera: Inocelliidae) . 41 C flickering , A. M. A new A canthoctenus (Araneae: Acanthoctenidae ) from Jamaica, W. I. 81 The Female of Lucarachne beebei Gertsch (Araneae: Symphytognathidae). 95 Christiansen, K. The Genus Pseudosinella (Collembola: Entomobryidae ) in Caves of the United States. 1 Crabill, R. E., Jr. On the True Nature of the Azygethidae (Chilopoda: Geophilomorpha). 76 Darlington, P. J ., Jr. Australian Carabid Beetles IV. List of Localities, 1956-1958. Ill Eisner, T. Defense Mechanisms of Arthropods. II. The Chemical and Mechanical Weapons of an Earwig. 62 Evans, H. E. A Study of Bembix u-scripta, a Crepuscular Digger Wasp. 45 MacLeod, E. G. The Immature Stages of Boriomyia fidelis (Banks) with Taxonomic Notes on the Affinities of the Genus Boriomyia (Neuroptera: Hemerobiidae) . 26 Werner, F. G. A Note on the Prey and a Nesting Site of Cerceris truncata Cameron. ( Hymenoptera : Sphecidae : Philanthinae) . 43 Wheeler, G. C. and J. Wheeler. Techniques for the Study of Ant Larvae. 87 INDEX TO SUBJECTS All new genera, new species and new names are printed in capital type. Acanthoctenidae, 81 Acanthoctenus , 81 A canthoctenus remotus, 81 Ant larvae, 87 Ants, 44, 87 Araneae, 81, 95 Arthropods, 62 Australian Carabid beetles IV. List of localities, 1956-1958, 111 Azygethidae, 76 Beetles, 111 Bembix u-scripta, 45 Boriomyia, 26 Boriomyia fidelis, 26 Carabid beetles, 111 Carboniferous insects, 98 Cerceris truncata, 43 Chilopoda, 76 Coleoptera, 111 Collembola, 1 Defense mechanisms of Arthropods. 11. The chemical and mechanical weapons of an earwig, 45 Digger wasps, 45 Earwig, 62 Entomobryidae, 1 Eurhopalothrix, 44 Eurhopalothrix bolaui, 44 Female of Inocellia pilicornis Carp. (Neuroptera: Inocelliidae) , 41 Female of Lucarachne beebei Gertsch (Araneae: Symphytognathidae) , 95 Genus Pseudosinella (Collembola: Entomobryidae) in caves of the United States, 1 Geophilomorpha, 76 Hemerobiidae, 26 Hymenoptera, 43, 44, 45, 87 Immature stages of Boriomyia fidelis (Banks) with taxonomic notes on the affinities of the genus Borio- myia (Neuroptera : Hemerobiidae) , 26 Inocellia pilicornis, 41 Inocelliidae, 41 Lucarachne beebei, 95 Neuroptera, 26, 41 New Acanthoctenus (Araneae: Ac- anthoctenidae) from Jamaica, W.I., 81 North American Carboniferous in- sects, 98 Note on the prey and a nesting site of Cerceris truncata Cameron. (Hymenoptera: Sphecidae: Philan- thinae), 43 Odonata, 71 On the true nature of the Azygethi- dae (Chilopoda : Geophilomorpha) , 76 Palae other ate s pcnnsylvanicus , 105 Paralogidae, 101 Paralogopsis longipes, 103 Paralogus aeschnoides, 102 Philanthinae, 43 Protodonata, 98 Pseudosinella, 1 Pseudosinella alba, 13 Pseudosinella argentea, 15 Pseudosinella boneti, 24 Pseudosinella dubia, 9 Pseudosinella duodecimpunctata, 8 Pseudosinella espana, 19 Pseudosinella folsomi, 18 Pseudosinella gisini, 14 Pseudosinella hirsuta, 22 Pseudosinella orba, 20 Pseudosinella petterseni, 15 Pseudosinella sexoculata, 12 Pseudosinella spinosa, 23 Rhopalothrix, 44 Rhopalothrix bolaui, 44 Sphecidae, 43 Studies on North American Carboni- ferous insects. I. The Protodonata, 98 Study of Bembix u-scripta, a crep- uscular digger wasp, 45 Symphytognathidae, 95 Techniques for the study of ant larvae, 87 Triassic odonate from Argentina, 71 TRIASSOTHEMIS, 71 TRIASSOTHEMIS MENDOZENSIS, 71 Troglosinella, 3 Troglosinella hirsuta, 22 Troglosinella sp'nosa, 23 Type species of the ant genus Eu- rhopalthrix, 44 Typus durhami, 98 Wasps, 45 PSYCHE Vol. 67 March-June, i960 Nos. 1-2 THE GENUS PSEUDOSINELLA (COLLEMBOLA, ENTOMOBRYIDAE) IN CAVES OF THE UNITED STATES1 By K. Christiansen Grinnell College, Grinnell, Iowa In 1934 Bonet listed two troglophile species of the genus Pseudo- sinelln, P. decipiens and P. sexoculata, from caves of the United States. In 1949 Delamare described two new troglobite species, P. hirsuta and P. spinosa, and placed them in a new genus Troglosinella. The present work is the result of the examination of more extensive collections from caves in the United States and has resulted in the addition of four more troglophile forms and the discovery of five new troglobite species. The work covered in this paper was made possible by a National Science Foundation Grant, No. G4563. Mr. Jerry Tecklin did much of the manual labor involved in preparation and he, Mr. James Hedges and Mr. George Darland, made a collecting trip through Missouri and northern Arkansas, recovering a good deal of critical material. Most of the remaining specimens were collected by Mr. and Mrs. Thomas Barr or Mr. Carl Krekeler. I wish to thank all of these people. Distribution Collections are relatively complete only from the region immediately around Tennessee and Kentucky; however, we have a sufficient scat- tering of material from other regions to be able to discern the probable outline of the distribution of the group. The species of this genus are more widespread in caves than those of the genus Sinella previously reported, but the troglobite forms are still largely limited to the southeast quadrant of the country. In the troglophile forms the ex- tensive invasion of the caves of Missouri and Arkansas is the most notable phenomenon. Generally speaking, the troglobite forms are severely restricted in distribution and, in cases of numerous collections, more or less continuous. The troglophile forms are much more wide- 'Published with the aid of a grant from the Museum of Comparative Zoology at Harvard College. I Smivhsuiwkn 1 § i§g 2 Psyche [March - June spread and occur in widely disjunct populations. As was true in Sinella , the majority of the collections of troglophile forms falls out- side of the areas occupied by the troglobite species. Biology Little accurate, detailed information is available concerning the biology of the members of this genus. The visible gut contents indicate that fungal hvphae and spores make up the major portion of the diet of most members. The incompleteness of the data on and collections of both cave and epigeic forms of this genus makes it difficult to classify the habitat of the various species. On a basis of present data we can place the forms described here into four groups: i) the definite troglo- GRAPH I Graph 1. Curves showing the differing ratios of the fourth antennal seg- ment to the cephalic diagonal in the cave species of U. S. Pseudo sinella. A includes: P. argentea, P. folsomi, P . duodecimpunctata , P. alba, and P. scxoeulata; B: P. dubia; C (a probable average of several curves): P. hirsuta ; D: P. gisini \ E: P. spinosa ; and F: P. boneti. bites — including P. spinosa, P. boneti, P. espana, P. hirsuta, and P. gisini; 2) the doubtful troglobites — P. dubia and P. orba ; 3) the doubtful troglophile — P. argentea ; and 4) the definite troglophiles — P. petterseni, P. alba, P. folsomi, P. duodecimpunctata, and P. sexoculata. 1960] Christiansen — Genus Pseudosinella 3 Systematic Account: General Discussion The genus Troglosinella was created by Delamare for the species hirsuta and spinosa. Examination of much more extensive material encompassing several new species makes it clear that the generic limits set up by Delamare are impractical. Two of the most basic character- istics used to separate the genus (the ringing of the fourth antennal segment, and the small non-lamellate teeth) are present in some forms of hirsuta and absent in others. Beyond this the reduced tooth structure of the unguis appears in a number of separate epigeic forms. The spines of the dens, also used to separate the genus, are found in only one species (P. spinosa ) and the “heavy hairs” found on the dorsum of the dens in hirsuta are found in large specimens throughout the whole of Pseudosinella. In view of this, and my failure to dis- cover any other practical way to separate the genus from Pseudosinella, I consider Troglosinella Delamare to be a synonym of Pseudosinella Schaffer. It is regrettable that this is unavoidable, since the species clustered around P. spinosa do represent an evolutionary unit. Ratios and Size As with almost all groups of Collembola, the size of the species varies considerably. It is extremely risky to determine the size of a particular species upon anything except large samples from a variety of localities. Time and again a whole sample will be made up of small individuals, or will consist exclusively of extra-large specimens. With all this in mind it is possible to break the cave species of Pseu- dosinella of the United States into three size ranges: small (averaging around I mm.) — P. espana (?), P. alba, P. folsomi, P. duodecim- punctata, and P. sexoculata ; medium (averaging between 2-3 mm.) — P. orba (?), P. boneti (?), P. dubia, P. gisini, P. argentea and P. hirsuta ; and finally the large (averaging around 4 mm.) — P. spinosa. The ratios of the various organs vary, but if we consider all the species, almost any organ ratio can be expressed as a straight line, with one or two notable exceptions. The only striking exceptions involving several species concern the first and second abdominal segments and the length of the antennae. The latter is illustrated in graph 1, which shows the most sensitive segment, the fourth, plotted against the cephalic diagonal. It can be seen that the most highly evolved cave forms develop progressively longer antennae. For most of the other organs a single straight line can express the growth changes and en- compass all species. Graph 2 shows a summary of such growth lines for organs of the various parts of the body. There are a few varia- 4 Psyche [March - June tions from the norm in several of these curves. The most striking is in connection with one population of P. hirsuta which has a longer fourth abdominal segment than is normal, and in all of the specimens examined of P. duodecim punctata which have the manubrium, longer than is normal in the remaining species. It is quite probable that more exhaustive analysis will point out more such exceptions; however, it is apparent that there is little difference among the various species in most organ ratios. The genus Pseudosine/la under any definition is a patently polyphy- letic group with characteristics largely determined by a particular ecology ; it can be distinguished from its ancestral genus Lepidocyrtus upon only one basis — the reduction in the number of eyes. Pseudosi- nella usually shows additional characteristics in the loss of pigment and the modification of the unguis structure, either through great elongation and reduction in the size and number of inner teeth and/or through the enlargement of the basal inner ungual teeth. Within the North American cave forms there are three and possibly four separate evolutionary lines involved. Only one of these, the group including Graph 2. Curves for five different organ ratios, each curve being a com- posite of all species. In the key below the ordinate is given first in each case. A: cephalic diagonal/second theoracic segment; B: cephalic diagonal/third tibiotarsus; C: manubrium/mucrodens ; D: cephalic diagonal/mucrodens ; E: cephalic diagonal/fourth abdominal segment. 1960] Chris tinmen Genus Pseudosinella 5 P. spinosa , has gone through extensive evolution within the caves. In spite of the diverse origins of the species of this genus the forms are normally quite similar. The thorax has a somewhat enlarged second segment and the mucro is bidentate with a basal spine. The chaeto- taxy is very uniform with macrochaetae on the head being commonly numerous only along the antennal bases, between the antennae, and on the anterior half of the ventral surface. The mesothorax has a series of rows along the anterior margin, and the lateral surface of the fourth segment and the last two segments have sparse coverings. Scattered macrochaetae and groupings of from two to four odd, short, feathered setae and one long filamentous ciliate seta occur on the third and fourth abdominal segments (See figure n). The scales are hyaline, finely striate and apically rounded. On the whole we have an artificial but readily separated genus. Key to the Cave Species of Pseudosinella of the United States I ) Empodial appendage with a clear wing-like tooth at the end of a separate outer lamella 2 lf) Empodial appendage with a small outer tooth or toothless 4 2) Mucro without basal spine P. espanci n. sp. 2') Mucro with basal spine 3 3) Unguis with median unpaired inner tooth P. petterseni 3') Unguis without median unpaired innner tooth P. folsomi 4) Dens with spines P. spinosa 4') Dens without spines 5 5) With eyes 6 5') Eyeless it 6) Two or fewer eyes per side 7 6') Three or more eyes per side1’ 9 7) All inner ungual teeth small 8 7') Basal, and sometimes all inner teeth large, two eyes per side P. alba 8) Tenent hair weakly clavate and/or median unpaired inner ungual tooth absent P . gisini n. sp. 8') Tenent hair acuminate, median unpaired inner ungual tooth present eyed forms of P. hirsuta 9) Three eyes per side on two separate eyepatches .... P. sexoculata 9') Some members of each population with more than three eyes per side, always on one eyepatch 10 10) All inner ungual teeth subequal, typically five eyes per side P. dubia n. sp. Occasional members of a population with four or more eyes may be eyeless. 6 Psyche [March - June i o') Basal inner ungual teeth clearly larger than others, typically six eyes per side P. duodecimpunctata ii ) Fourth antennal segment more than two times as long as cephalic diagonal P. ho net i n. sp. n') Fourth antennal segment less than one and seven-tenths times as long as cephalic diagonal 12 12) Distance from distal base of largest inner ungual tooth to base of unpaired inner tooth less than half that from unpaired tooth to apex of unguis (See figure 44), macrochaetae present in median field of dorsum of head P. orha n. sp. 12') Distance from largest basal inner tooth to median unpaired tooth at least half as great as distance from apical tooth to apex of unguis, macrochaetae absent from median field of dorsum of head 13 13) One or both basal inner ungual teeth large and prominent (See figure 31 ), ratio of fourth antennal segment to cephalic diagonal less than .8 P. ar gen tea 13') Both basal inner ungual teeth small (See figure 47), and/or ratio of fourth antennal segment to cephalic diagonal greater than .8 eyeless forms of P. hirsuta Explanation of Plate 1 Figures 1-5: Pseudosinella duodecimpunctata. 1. Habitus, specimen from New River Cave, Giles Co., Va.; setae and pigment omitted; 19 X. 2. Hind claw, same specimen; 900 X. 3. Mucro, same locality; 900 X. 4. Apical organs, third antennal segments two different specimens, same locality; 900 X. 5. Eyes and eye patch, same locality; 900 X. Figures 6-11: P. dubia n. sp. 6. Habitus, specimen from Granny Dean Cave, Washington Co., Ark.; setae and pigment omitted; 18 X. 7. Apical organ, third antennal segment, same specimen; 900 X. 8. Eye patch right side, same specimen; 250 X. 9. Mucro, same specimen; 500 X. 10. Hind claw, same specimen; 500 X. 11. Typical setae association of fourth abdominal segment, paratype; 900 X. Figures 12-16: P. sexoculata. 12. Habitus, specimen from Carlsbad Caverns, N. Mex., setae omitted, 30 X. 13. Mucro, same specimen, 900 X. 14. Apical organ third antennal segment, same specimen; 900 X. 15. Hind claw, same specimen; 900 X. 16. closeup of head, same specimen, A.-interantennal setae, B. -antennal base setae, C. -median field setae; 150 X. Figures 17-21: P. alba. 17. Habitus, specimen from Niagra Cave, Fillmore Co., Minn.; 30 X. 18. Hind claw, same specimen; 900 X. 19. Apical organ, third antennal segment, same specimen; 900 X. 20. Mucro, same specimen; 900 X. 21. Right eye patch, specimen from Mystery Cave, Fillmore Co., Minn. ; 300 X. Figures 22-24: P. folsomi. 22. Apical organ, third antennal segment, speci- men from Limberlost Valley Cave, Newton Co., Mo.; 900 X. 23. Hind claw, same specimen; 900 X. 24. Mucro, epigeic specimen from Harahan, La.; 500 X. Psyche, 1960 Vol. 67, Plate 1 Christiansen — Pseudosinella 8 Psyche [March - June Descriptions of Species Pseudosinella duodecimpunctata Denis Figures 1-5 Pseudosinella duodecimpunctata Dennis 1931, Mem. Soc. Ent. Ital. 10:8 2. P. (Pseudosinella) duodecimoculata Bonet 1931, Mem. Soc. Esph. N.H. 74:3 24-6. Facies typical of genus Lepidocyrtus. Background color yellowish white, with scattered blue pigment, particularly heavy along the sides of the thorax and head. Head broadly oval in shape with a clear V- shaped dorsal area marked off by pigment ; six eyes per side on trape- zoidal eye patches. Antennae 1.2- 1.4 times as long as cephalic diagonal with third antennal segment in the form of a truncate cone clearly shorter than the second segment; apical organ of third segment with two irregular oval pegs in separate deep folds. Second thoracic slightly humped. Fourth abdominal segment about four times as long as third. Unguis with a clear external tooth and three inner teeth; basalmost of these usually salient and larger than remainder. Em- podial appendage lanceolate, with one or more minute external teeth. Tenent hair curved and clearly clavate. Mucro with apical tooth weakly upturned and longer than anteapical, basal spine attaining the apex of the anteapical tooth. Anterior macrochaetae as follows: on dorsum of head a group of four interantennal setae near the anterior margin and a curved row of seven setae along each antennal base, the anterior three being distinctly smaller; ventral surface with a few scattered setae near the anterior margin of the median and lateral areas. Second thoracic segment has a double row along the anterior margin. Type locality: Buco de Piombo, Italy. Distribution: widespread in southern Europe in caves and epigeic. In North America the species has been taken in Florida, Massachusetts, and North Carolina as an epigeic form. In caves of the United States it has been taken from New River Cave, Giles County, Virginia; Wind Cave, Jackson County, Kentucky; Old Spanish Cave, Stone County, Missouri; Panther Cave, Newton County Missouri; and Foster’s Cave, Mont- gomery County, Tennessee. D iscussion The most striking variations seen in this troglophile species are in the number of eyes and amount of pigment. Most specimens have the twelve eyes characteristic of the species; but forms with ten eyes 1960] Christiansen — Genus Pseudosinella 9 are common ; rarely, specimens appear with fewer, as can be seen in those from Tennessee which have from none to six per side. The tibiotarsus of this population is slightly longer than is normal for the species, but they are otherwise similar to forms seen elsewhere. This would appear to be an example of cave evolution proceeding within a population and would be worth additional study. Pigment may be uniformly distributed over the whole body or limited to the head region. The apical organ of the third segment varies from the condi- tion described to two subcylindrical pegs. The identity of this species with the European specimens is still in doubt as no comparisons could be made; however, there appears to be good agreement between the descriptions and the specimens at hand. This species may very well be the same as P. collina , which was de- scribed by Wray (1952) as having only two internal ungual teeth; but until this can be checked the two species must be considered sepa- rate. Pseudosinella dubia, new species Figures 6-1 1 Facies typical of genus. Background color white with an overall scattering of blue pigment, particularly dark on dorsum of head where it forms a diamond shaped mark between the eye patches. Head broadly oval; eyepatches elongate trapezoids, each bearing five eyes in two groups, an anterior group of three and a posterior group of two. Antennae about one and one half times as long as the cephalic diagonal ; first three segments subcylindrical and fourth segment fusi- form. Fourth segment dorsally with numerous blunted smooth setae of several different sizes and shapes; apical organ of third segment with two irregular flattened elliptical rods in a fold, and about fifteen additional blunt curved setae scattered over the inner ventral surface of the segment. Second thoracic segment markedly humped forcing the head into a slightly hvpognathous position. Fourth abdominal segment about five times as long as third. Unguis with minute basal external tooth and three small internal teeth, with basal-most inner tooth on a level and frequently appearing as one under low magnifica- tion. Empodial appendage lanceolate, with a small but clear external teeth, exceeding the level of the unpaired internal ungual tooth. Ten- ent hair weakly clavate. Mucro with apical tooth about twice as long as subapical, and markedly upturned ; basal spine exceeding the apex of subapical tooth. Anterior macrochaetae as follows: on dorsum of head four interantennal setae, the posterior pair closer together, and IO Psyche [March - June a curved row of eleven setae around each antenna base ; ventral surface with a few scattered setae on the anterior median portion. Type locality: Devils Den Kitchen Cave, Washington County, Arkansas, IX-9-’59, Tecklin, Hedges and Darland coll. Also taken from Granny Dean and Devils Den Caves, Washington Countv. Arkansas. Discussion The number of eyes varies considerably, most specimens having five per side. However, many have four per side ; in such cases it is the posteriormost eye of the anterior three which is missing, and the position of this eye is variable even when it is present. The tenent hair varies from acuminate to markedly clavate. The taxonomic position, name and limits of this species are all moot. The species is clearly distinguished from the form called here P. duodccim punctata. The antennae of duhia are longer, and the eye number typically different. The shape of the eye patch and distribution of the eyes are also different, as is the structure of both mucro and unguis. The question of the relationship between this form and Wray’s collina or Guthrie’s Lepidocyrtus decemoculatus is less easily settled. The present species appears to differ from collina in (i) the number of eyes (six vs. five per side typically) ; (2) the relative lengths of abdominal segments three and four ( collina one: four, d.ubia one: five + ) ; (3) antennal ratios; and (4) the comparison of lengths of the manubrium and dens. In addition, Wray’s figures show the basal spine of the mucro not reaching the anteapical tooth and the unguis Explanation of Plate 2 Figures 25-30: P. gisini n. sp. 25. Habitus, specimen from Higgenbotham Cave, Greenbrier Co., W. Va., setae and pigment omitted; 12 X. 26. Hind claw, paratvpe; 350 X. 27. Hind unguis, specimen from McFerrin Cave, Greenbrier Co., W. Va.; 250 X. 28. Mucro, same specimen; 250 X. 29. Dor- sal setae and eyes right side, paratype, Greenbrier Co., W. Va.; A.-inter- antennals, B. -antennal base setae; 120 X. 30. Apical organ, third antennal segment, specimen from McClung Cave, Greenbrier Co., W. Va.; 350 X. Figures 31-37: P. argentea. 31. Typical hind unguis, specimen from Eli Reed Cave, Larue Co., Ky. ; 350 X. 32. Hind claw showing unusually large basal tooth, specimen from Rankin Cave, Jefferson Co., Mo.; 350 X. 33. Hind unguis showing unusually small basal teeth, specimen from Crownover Saltpeter Cave, Franklin Co., Tenn.; 350 X. 34. Habitus, specimen from Mammoth Cave, Ky., setae omitted; 17 X. 35. Apical organ, third antennal segment, specimen from Rankin Cave, Jefferson Co., Mo.; 900 X. 36. Same organ, specimen from Sparkman Cave, White Co., Tenn.; 900 X. 37. Mucro, same specimen ; 500 X. Figures 38-42: paratypes of P. espana n. sp. 38. Habitus, setae omitted; 30 X. 39. Mucro; 900 X. 40. Apical organ, third antennal segment; 900 X. 41. Base of hind claw; 900 X. 42. Claw smaller specimen; 900 X. Psyche, 1960 Vol. 67, Plate 2 Christiansen — Pseudosinella 12 Psyche [March - June with two inner teeth. Both conditions differ in dubia. Lepidocyrtus decemoculatus is very poorly described, but every major described featuie except the eye number would appear to indicate a separate species. The distribution of the eyes, the shape of the eyepatch, the hinted antennal ratios, and ratio of manubrium to dens all are differ- ent from those seen in dubia. In any case, I feel that it is impossible to fit the present form into the desci lptions of either species at this time. It is quite possible that close examination will show that this form is in fact synonymous with one or both of the species mentioned above. Pseudosinella scxoculata Schott Figures 12-16 Pseudosinella scxoculata Schott 1902, Bib. Kong. Sv. Vet.-Akad; Handl. 28 : 34-5. Lepidocyrtus sexoculatus Guthrie 1903, Publ. Geol. Nat. Hist. Survey Minn. Zool. Series 4 : S6-7. P ( seudosinella) scxoculata Bonet 1934, Arch. Zool. Exp. Gen. 76: 370. Facies typical of genus Lepidocyrtus. Background color dull yellow, pigment limited to eyepatches. Head circular, eyes three per side on two separate eye patches, two eyes in front and one behind. Antennae with fourth segment fusiform, second and third segments truncately conical and first segment subcylindrical ; third segment strikingly shorter than the second and with an apical organ consisting of two basally constricted oval pegs in a deep fold. Second thoracic segment not strikingly humped, so that the head is prognathous. Fourth abdom- inal segment about three times as long as the third segment. Unguis without external teeth, the basal pair arising at different levels. Empodial appendage lanceolate with outer margin serrate. Tenent hair large and clearly clavate. Mucro with apical tooth slightly larger than anteapical and markedly upturned at its apex; basal spine exceed- ing apex of anteapical tooth. Anterior macrochaetae as follows: dorsum of head with a curved row of six setae along each antennal base and a single seta between the two eyepatches on either side; four interantennals, more posterior than normal ; a medium pair of setae between and slightly in back of the two posterior eye patches; ventral surface of head with scattered setae along the anterior one-fourth. Second thoracic with a double row of setae along the anterior margin. Type locality: epigeic greenhouses, Linkoping, Sweden and Rosen- dal, Norway. 1960] Christiansen — Genus Pseudosinella 13 Distribution: a common epigeic form on. the Pacific coast. Cave collections in the United States: Reids Cave, Fayette County, Ken- tucky; and Carlsbad Caverns, Eddy County, New Mexico. Discussion This fo rm appears to be a rare troglophile in caves in the United States. Further exploration and collecting in western caves may show it to be more common than presently appears. Pseudosinella alba (Packard) Figures 17-21 Lepidoeyrtus albus Packard 1873, Peabody Acad. Sci. 5th Ann. Rept: 37- Sira ( Pseudosinella ) alba Schaffer 1900, Jahib. Ver. Vaterl. Natur. 56 : 269. Pseudosinella alba Borner 1901, Zool. Anz. 24: 707. Facies typical of genus Lepidoeyrtus. Background color dull yel- low white with scattered bluish or brownish pigment. Eyes two per side on square black pigment patches. Fourth antennal segment fusi- form, third segment truncately conical and remainder subcylindrical ; third segment noticeably shorter than second and with apical organ of two short subcylindrical pegs. Second thoracic segment slightly en- larged, with head prognathous. Fourth abdominal segment about four times as long as third. Unguis with three inner teeth with basal- most tooth being the largest. The empodial appendage is lanceolate with a small but clear external tooth. Tenent hair clavate. Mucro with apical tooth about twice as long as anteapical and not strikingly upturned ; basal spine barely attaining apex of anteapical tooth. Anterior macrochaetae as follows : on the dorsum of head a group of four setae along the base of each antenna, and a group of four interan- tennal setae forming the corners of a narrow rectangle; in the center are four more setae marking the sides of a triangle which has its apex just anterior to the anterior pair. Ventral surface of head sparsely covered with setae over the anterior median portion. Anterior margin of second thoracic segment with a double row of setae. Type locality: epigeic but uncertain, probably Massachusetts. A common epigeic form in the United States known from Mystery and Niagra Caves in Fillmore County, Minnestota and Christian Cave in Robertson County, Tennessee. Discussion I have examined and measured a considerable number of epigeic forms of this species and can find no significant differences between 14 Psyche [March - June these and the cave forms. The coloration is the most variable charac- teristic; some specimens are white save for the eyespots, while all others are almost totally blue. Pseudosinella gisini , new species Figures 25-30 Facies typical of genus. Color white with a scattering of blue pig- ment particularly clear upon the dorsum of the head. Head elongate oval with two eyes per side located upon a single dark eyepatch. Antennae about three and one half times as long as cephalic diagonal with all segments subcylindrical in shape; fourth antennal segment without apical cone and showing definite signs of ringing; third seg- ment with apical organ of two curved, narrow, paddle-shaped setae with supporting rods along one margin. On the opposite side of the apex of third segment are five blunt finger-like setae. Second thoracic segment not humped. Fourth abdominal segment about four times as long as third. Unguis long and curved with external and lateral teeth minute and three inner teeth, basal pair small but clear and on a level, median unpaired tooth minute, visible only under highest magnifica- tion. Empodial appendage with slight basal internal swelling. Tenent hair finely clavate. Mucro with teeth subequal, apical tooth not sharp- ly upturned, basal spine heavy, reaching apex of anteapical tooth. An- terior macrochaetae as follows: dorsum of head with four interanten- nal setae forming the corners of a rectangle, and a row of eight setae forming a straight line along each antenna base; ventral surface with scattered setae over the anterior half of the median and lateral sur- faces. Second thoracic segment with two to three rows of setae along the anterior margin and a single row along the lateral margin. Type locality: Foxhole Cave no. 1, Greenbrier County, West Vir- ginia, VIII- 19 ’58. Barr coll. Also known from eight additional caves, all in Greenbrier County. Discussion The most prominent variation seen in the species concerns the median unpaired tooth of the unguis which may be entirely absent. In such forms the tenent hair is very weakly clavate so that this is visible only under highest magnification and with phase contrast. Although the typical form of this species is strikingly distinct from the typical form of P. hirsuta there are some specimens of the latter species which approach gisini in one or more characteristics (see discussion under P. hirsuta) ; however, the two can always be distinguished on a basis of the cephalic chaetotaxy and the tenent hair. 1960] Christiansen — Genus Pseudosinella 15 Pseudosinella petterseni Borner Pseudosinella petterseni Borner 1901, Zool. Anz. 24: 707-8. Facies typical of genus. Color silvery white without trace of pig- ment. Head circular and eyeless. Antennae with fourth segment fusi- form, third segment truncately conical and remainder subcylindrical ; apex of fourth segment rounded ; apical organ of third segment with two subcylindrical pegs. Second thoracic segment not humped and the head prognathous at rest. Fourth abdominal segment about three and one half times as long as the third segment. Unguis with a small external tooth, and three well developed internal teeth; the basal pair not on a level and the basalmost one considerably larger than the re- mainder. Empodial appendage with a large outer wing tooth on a distinct lamella. Tenent hair strikingly clavate. Mucro with apical tooth twice as long as anteapical ; basal spine exceeding the apex of anteapical tooth. Anterior macrochaetae as follows: dorsum of head with a row of six setae along each antenna base, the setae in each row being alternately large and small, the posteriormost seta of each row is displaced inward; interantennal setae are concentrated near the an- terior margin of the head so that only two are clearly visible; ventral surface with scattered large setae on anterior one third of surface. Anterior margin of second thoracic segment with from two to four rows of setae. Type locality: epigeic, Frauenberg bei Marburg, Germany. Distribution: one of the commonest epigeic species in the United States; known in caves from Kendall and Burnett Counties, Texas, and Linn County, Iowa. Discussion The specimens seen from the Texas caves differ slightly from the epigeic forms seen in most of the country ; in the former the basal ungual tooth is smaller, the tenent hair clavate and the basal spine of the mucro shorter ; however, these variations are probably geograph- ical since epigeic specimens seen from New Mexico indicated similar modifications. Further collections from western caves will probably show this to be a much more common cave inhabitant than is presently indicated. Pseudosinella argentea Folsom Figures 31-37 Pseudosinella argentea Folsom 1902, Psyche 9: 366. P (seudosinella) dccipiens (?) Bonet 1934, Arch. Zool. Exp. Gen. 76: 370-3. i6 Psyche [March - J une Facies typical of genus. Color white with scattered blue pigment particularly upon head. Head almost round when seen from above, often with traces of pigmentation on eyespot region ; eyeless ; antennae with third segment sub-conical in shape; fourth antennal segment much shorter than cephalic diagonal and with apex rounded, without apical cone. Apical organ of third segment of two strongly curved blunt pegs with an additional pointed peg off to one side. Thorax not humped and head prognathous. Fourth abdominal segment about four times as long as the third segment. Unguis with one or both basal teeth enlarged and a single median tooth prominent and projecting. Empodial appendage lanceolate with a very small external tooth. Tenent hair weakly clavate. Mucronal teeth subequal and basal spine attaining the apex of the median tooth. Anterior macrochaetae as follows: dorsum of head with four interantennal setae arranged in a narrow rectangle and with a small seta located in the center of this rectangle. Antennal bases with seven setae arranged 6-1 in a straight line along the base. Venter of head with a few scattered setae near the anterior margin. Second thoracic segment with three to four rows along the anterior margin and a single row of smaller setae along the lateral margin. Type locality: a grave, Washington, D. C. Range: known from caves in Missouri, Central Tennessee, and single localities in Ken- tucky, Virginia, Arkansas, Georgia and Washington, D. C. Discussion This species is almost as variable as P. hirsuta «nd in some forms the basic structure is very similar. The two species can always be sep- Explanation of Plate 3 Figures 43-46: holotype of P. orha n. sp. 43. Habitus, setae omitted; 30 X. 44. Hind claw; 500 X. 45. Apical organ, third antennal segment; 900 X. 46. Mucro ; 500 X. Figures 47-58: P. hirsuta. 47. Hind claw, specimen from Ward Cave, Bedford Co., Tenn.; 350 X. 48. Unguis, another specimen same locality; 350 X. 49. Hind unguis, specimen from Mill Creek Cave, Davidson Co., Tenn.; 250 X. 50. Apical organ, third antennal segment, same specimen as fig. 47; 900 X. 51. Habitus, topotype ; 18 X. 52. Mucro, same specimen as fig. 47; 900 X. 53. Mucro, topotype; 900 X. 54. Mucro, same specimen as fig. 49; 900 X. 55. Apical cone fourth antennal segment, specimen from Piper Cave, Smith Co., Tenn.; 900 X. 56. Antennal base setae, same specimen; 250 X. 57. Thoracic profile, specimen from Gassaway Cave, Metcalfe Co., Ky. ; 30 X. 58. Thoracic profile, specimen from Pratt Cave, Pickett Co., Tenn.; 30 X. Figures 59-61: P. spinosa. 59. Mucro, specimen from Jared Hollow Cave, Putnam Co.. Tenn.; 900 X. 60. Apical organ, third antennal segment, speci- men from Payne Saltpeter Cave, Grundy Co., Tenn.; 900 X. 61. Hind claw, same specimen; 250 X. Psyche, 1960 Vol. 67, Plate 3 Christiansen — Pseudosinella i8 Psyche [March - June arated on a basis of the cephalic chaetotaxy and the apex of the an- tenna. The variations of the unguis structure are most striking. Typically the median tooth is large, but it may be small (see figures) ; also the basal teeth are normally staggered and of different sizes, but they may be of equal size and/or at the same level. The external tooth is usually present but may be folded and virtually invisible. A small apical tooth may be present. The tenent hair varies from strik- ingly clavate to acuminate. Some forms are totally without pig- ment while others are clearly colored. The cephalic diagonal, fourth antennal segment and head also vary considerably in size. The an- tennal base setae also vary with size (as is normal for the genus) but the huge majority of specimens examined had the number noted. First examination of this material led to the belief that two or three species were involved ; but, with larger series of specimens from many localities and careful analysis of the variation, it became ap- parent that no clear boundaries could be drawn within the whole group and that the combinations of characteristics were virtually at random. It appears that some of the variation is geographic in nature (example: large median tooth more common in northern populations) but an analysis of this must await further and more complete collections. Specimens taken from caves in Virginia and identified by Bonet as P. decipiens (1934) are almost certainly argentea. It is quite possible that P. argentea is in fact synonymous with P . decipiens. The unguis in particular appears to have much the similar sort of variation in the European forms as in argentea. The figures for this species given by Denis, Bonet and Gisin would appear to indicate considerable varia- tion; however, there are certain striking differences. First, in argentea a common form has both teeth arising at the same level, and this ap- parently is never true with decipiens. Second, Denis indicates the api- cal organ of the third antennal segment in decipiens as consisting of two cylindrical, straight pegs, whereas in argentea they consist of two basically constricted, strongly curved elliptical pegs. The fourth an- tennal segment is relatively longer in argentea than Bonet indicates for decipiens, and the fourth abdominal segment is from four to five times as long as the third rather than 3 to 3.5 times longer as indicated by Bonet for decipiens. In view of all of these I feel it wiser to maintain argentea until such time as a careful comparative study can be made between this form and decipiens. Pseudosinella folsomi Denis Figures 22-24 Pseudosinella folsomi Denis 1931, Mitt. Mus. Hamburg 44: 226-8. 1960] Christiansen — Genus Pseudosinella 19 Facies typical of genus. Color white without vestige of pigment. Head round, eyeless. All antennal segments subcylindrical ; apex of fourth segment blunt ; apical organ of third segment with two large elliptical pegs, constricted basally. Second thoracic segment slightly humped, head more or less prognathous. Fourth abdominal segment about three times as long as third. Unguis with only two strong basal internal teeth, arising at about same level and with one greatly enlarged. Empodial appendage with inner margin slightly truncate, and outer margin equipped with a prominent tooth on a distinct lamella. Tenent hair short and clavate. Mucro with apical tooth slightly longer than anteapical ; basal spine not attaining apex of anteapical. Anterior macrochaetae as follows : dorsum of head with a group of five at each antenna base and four interantennals. Another pair of setae can be seen near the lateral margins of the middle of the dorsum ; ventral surface with numerous scattered setae on the anterior third of the lateral and median portions. Anterior margin of the second thoracic segment with four to five rows of setae and a single row along the lateral margins. Type locality: epigeic “Station No. 36,” Massachusetts. Also re- corded as an epigeic form from several places in the vicinity of Boston and New Orleans. Cave distribution: known only from Limberlost Valley Cave, Newton County, Missouri. Discussion Recent collections from South America would seem to indicate that this species is widespread in Chile at least. I he fact that the epigeic collections in the United States have been limited to the vicinity of major ports makes it seem probable that this is an imported form here. Its one collection in a cave was in association with another troglophile species, Sinella caeca. Further collections will have to be made before anything can be said about the significance of this in- vasion. Pseudosinella espana, new species Figures 38-42 Facies typical of genus. Color white without trace of pigment. Head oval, eyeless. Antennae one and one half times as long as cephalic diagonal, with the first three segments subcylindrical and fourth segment ellipsoidal ; apical organ of third segment with two paddle-shaped setae having central supporting rods. Second thoracic segment not strikingly humped. Fourth abdominal segment slightly more than three times as long as third. Unguis narrow and sickle- 20 Psyche [March - June shaped with only two large basal internal teeth. Empodial appendage with a basal internal swelling and a large external tooth upon a dis- tinct lamella. Tenent hair small and acuminate. Mucro with apical tooth gradually curved and more than twice as long as anteapical ; basal spine absent. Anterior macrochaetae as follows: dorsum of head with five interantennal setae and a row of six setae near each antenna base and a pair near the center of the dorsal field ; a diagonal row of three and group of four setae near each latero-posterior margin of the dorsum; ventral surface with two mid-lateral patches of densely packed setae. Second thoracic segment with three to four rows of setae along the anterior margin and a single row along the lateral margins. Type locality: Old Spanish Cave, Stone County, Missouri, VI ’59, Tecklin, Darland, and Hedges coll. Also known from Coleman’s Cave, Montgomery County, Tennessee. It is probably widespread but uncollected in caves between these two points. Discussion This species is the only Nearctic member of the genus to lack a basal spine. Except for this remarkable feature it is very similar to P. folsomi although it shows a number of cave adaptions not found in th is last species. Pseudosinella orba, new species Figures 43-46 Facies typical of genus. Color white without trace of pigment. Head circular, eyeless. Antennae about twice as long as cephalic diagonal, all segments slightly ellipsoidal; second and third segments subequal in size; apical organ of third segment with two blunt oval pegs. Second thoracic segment slightly humped. Fourth abdominal segment about three times as long as third. Unguis broad, apically slightly hooked, basal internal teeth large, subequal, and arising well above the base of the claw; median unpaired tooth prominent and about at mid-level of unguis; a pair of small lateral teeth are near the base. Empodial appendage lanceolate, with external margin serrate and basally expanded. Tenent hair slender and acuminate. Explanation of Plate 4 Figure 62: P. spinosa, Habitus, specimen from Payne Saltpeter Cave, Grundy Co., Tenn., setae omitted; 18 X. Figures 63-65: P. honeti n. sp. 63. Habitus, holotvpe ; 18 X. 64. Mucro, holotype ; 350 X. 65. Hind claw, para- type; 250 X. Figure 66: Map showing known cave locations of six troglophile species of Pseudosinella in the U. S. Figure 67: Map showing cave distribu- tion of seven troglobite species of Pseudosinella in the U. S. Psyche, 1960 Vol. 67, Plate 4 Christiansen — Pseudosinella 22 Psyche [March - June Mucro with apical tooth much larger than anteapical; basal spine slender but reaching the apex of the anteapical tooth. Anterior macrochaetae as follows: dorsum of head with an interantennal pair, and then in the anterior half of the median field six more arranged along the sides of a triangle with its apex near the interantennals ; a straight row of eight setae along each antenna base; ventral surface with scattered setae along the anterior third. Second thoracic seg- ment with two rows along the anterior margin and a transverse row of eight setae along the posterior margin. Type locality: Morril Cave, Sullivan County, Tennessee. X ’56, T. Barr coll. Also taken from Hamilton Cave, Bland County, Vir- ginia. Discussion This striking species is quite unique in the chaetotaxy of the head and the structure of the unguis. In both characteristics it appears to be related to P. alba and may well represent a cave derivative of this species. The fact that the modifications for cave life are relatively slight makes it entirely possible that this is in fact a troglophile form which is so far undescribed from epigeic habitats. Pseudosinella hirsuta Delamare (new combination) Figures 47-58 Troglosinella hirsuta Delamare 1949, Notes Biosp’e., 4: 12 1-2. Facies typical of genus. Background color yellowish white with slight touches of blue or reddish (?) pigment, particularly in the region of the antennal bases. Head elliptical, normally without eyes. Antennae elongate with all segments subcylindrical ; fourth segment usually showing signs of ringing with a definite small conical projection at the apex; apical organ of third segment with two curved elliptical paddles, constricted basally. Second thoracic segment slightly humped. Fourth abdominal segment about six times as long as the third. Unguis elongate with a small external and three small internal teeth, the basal pair being on the same level. Empodial appendage narrow and lanceolate with a small external tooth. Tenent hair small and acuminate. Mucro with apical tooth about twice as long as anteapical and weakly upturned at the apex; basal spine just reaching the apex of the anteapical tooth. Anterior macrochaetae as follows: dorsum of head with an interantennal group marking the corners of a narrow rectangle; a row of nine setae along each antenna base with the posterior one or two in each row displaced inward ; ventral surface 1960] Christiansen — Genus Pseudosinella 2 3 with scattered setae along the anterior half. Two to three rows of setae along the anterior margin of the second thoracic segment. Type locality: Tennessee caverns, Tennessee, H. Henrot coll. Distribution: major distribution in south-central Kentucky, central Tennessee and northeast Alabama ; isolated populations in eastern Tennessee and Kentucky. Discussion This species is the most abundant and most variable troglobite form of the genus in North America. No single description can apply to all of the available specimens, but the characteristics listed above apply to about 6o% of the specimens and with one or another major alteration would fit all save about i% of the specimens seen. The variation is extensive and can be seen in virtually all characteristics so that some specimens resemble P. argentea while others are superfi- cially similar to P. gisini) indeed single specimens are often hard to place. Analysis of the variable characteristics shows that they are definitely geographically determined ; however, since there is no sim- ilarity among the distribution of the various characteristics, the sub- species concept is clearly not applicable. In general there are six characteristics which vary strikingly and these and their limits of variation are listed below: A) Second thoracic segment, strongly humped to virtually fiat. B) Unguis, condition seen in argentea to condition seen in gisini. C) Empodial appendage, lanceolate to clearly basally expanded. D) Ratio of fourth antennal segment to cephalic diagonal, 3:6- 5U-5- E) Mucro, anteapical tooth middle to near apical tooth. F) Eyes, eyeless to two eyes per side. G) Pigment, totally white to head and thorax medium blue. All in all the majority of the specimens are very similar to the form in the description and no one specimen carries very many of the variant characteristics. Pseudotisinella spinosa Delamare (new combination) Figures 59-62 Troglosinella spinosa Delamare 1949, Notes Biosp’e., 4: 122-4. Body elongate with attenuate appendages. Background color dull yellowish white with occasional flecks of reddish ( ?) pigment around the bases of the antennae. Head elliptical, strikingly longer than broad. Antennae with all segments subcylindrical, and fourth seg- ment showing signs of ringing and subsegmentation ; apex of antenna 24 Psyche [March - June with a small apical cone; apical organ of third segment with two flattened, fusiform, basally constricted paddles, in separate shallow folds. Second thoracic segment strongly humped forcing head into an opisthognathous position. Fourth abdominal segment about seven times as long as third. Unguis strikingly elongate, without external teeth but with three very small internal teeth, the basal pair being on a level and slightly smaller than the median unpaired tooth. Empodial appendage lanceolate, slender and with a striking internal basal ex- pansion. Tenent hair small and acuminate. Mucro with anteapical tooth displaced toward apex; teeth subequal in size; basal spine not attaining level of anteapical tooth. Dens with one or two rows of heavy finely ciliate spines along anterior two thirds. Anterior macro- chaetae as follows : dorsum of head with an interantennal group of five setae; a curved row of nine setae along each antennal base; ven- tral surface covered with numerous setae. Second thoracic segment with two to three rows of setae along anterior margin and a single row along the lateral margins. Type locality: Alladin’s Cave, Madison County, Alabama, H. Henrot coll. Distribution : limited to central Tennessee and north- east Alabama. Abundant within these limits. Discussion This is by far the most stable of the common troglobite species of this genus. The only striking variation is seen in the mucro, which has occasional populations with the anteapical tooth much more basal than is normal. Other variations are minor; for example, some forms are entirely white while others have a scattering of pigment over the whole body. The inner ungual teeth are often so minute as to be invisible under low magnifications. The dental spines are often difficult to observe. Pseudosinella boneti, new species Figures 63-65 Body elongate, appendages very attenuate. Color white without trace of pigmentation. Head oval without eyes. Antennae four or more times as long as head with all antennal segments subcylindrical ; apical organ of third antennal segment with two paddle-shaped setae having the supporting rod along one margin. Second thoracic seg- ment greatly enlarged, forcing the head into a hypognathous or opis- thognathous position. First abdominal segment strikingly compressed; fourth abdominal segment almost seven times as long as third. Unguis extremely thin and elongate having only a single pair of in- 1960] Christiansen — Genus Pseudosinella 2 5 ternal basal teeth. Empodial appendage thin and basally swollen. Tenent hair minute and acuminate. Dens without spines. Mucro with apical and anteapical teeth close together and near apex, sub- equal in size , basal spine not attaining level of anteapical tooth. Antei 101 maciochaetae as follows! dorsum of head with seven inter- antennal setae and a row of eight setae along each antenna base; ven- tral surface with numerous densely packed setae along median and lateral area. Anterior margin of second thoracic segment with two to three rows of densely packed setae. Type locality: Pomps Cave, White County, Tennessee, III ’57 T. Bail coll. Also taken from Dairyhouse Cave in the same county; Wolf Cave, Morgan County, Alabama; and Wind Cave, Pulaski County, Kentucky. Discussion This well marked species represents the apogee of cave evolution in the Nearctic Pseudosinella. The series available is limited and little variation has been seen. In some forms the ungual teeth are entirely absent, and in the specimens from Alabama and Kentucky the median tooth of the mucro is less apical than in the Tennessee specimens. The specimens from Kentucky sometimes have a very small median internal ungual tooth. Literature Cited Bonet, M. F. 1931. Estudios sobre coilembolos cavernicolas con especial referenda a los de la fauna Espanola. Mem. Soc. Hist. Esp. Nat. 14: 221-403. 1934. Biospeleogica No. XL. Campagne speleoligique de C. Bolivar et R. Jeannel dans 1’Amerique du Nord. 10 Collemboles. Arch. Zool. Exptl. et Gen. 76: 361-377. Borner, C. 1901. Neue Collembolen Formen und zur Nomenclatur der Col- lembolen Lubbock. Zool. Anz. 24: 696-712. Delamare Deboutteville, Cl. 1949. Collemboles Cavernicoles du Tennessee et de L’Alabama. Notes Biospeleoligiques. 4: 117-124. Denis, J. R. 1931. Sur la Faune itasienne des Collemboles. IV Notes pre- liminares de Collemboles de Caverne Italienne. Mem. Soc. Ent. Ital. 10 (1) : 80-85. Guthrie, J, E. 1903. The Collembola of Minnesota. Rep. Geol. Nat. Hist. Surv. Minn. Zool. Ser. (4): 1-110. Packard, A. S. 1873. Synopsis of the Thysanura of Essex county with a description of a few extralimital forms. Ann. Rep. Peabody Acad. Sci. 1873: 23. Schaffer, C. 1897. Hamburger Magalhaenische Sammelreise. Apterygoten 8: 1-48. Schott, F. 1902. Etudes sur les Collemboles du Nord. Bih. Svenska Vet. Akad. Handl. 28 (2) : 1-48. Wray, D. L. 1952. Some New North American Collembola. Bull. Brook. Ent. Soc. 47 (4) : 95-106. THE IMMATURE STAGES OF BORIOMYIA FIDELIS (BANKS) WITH TAXONOMIC NOTES ON THE AFFINITIES OF THE GENUS BORIOMYIA ( NEUROPTERA :HEMEROBIIDAE ) By Ellis G. MacLeod University of Maryland College Park, Maryland Introduction The genus Boriomyia Banks, 1904 ( sensu Killington, 1937, Car- penter, 1940; nec Banks, 1905, Tjeder, 1941, et seq.) is known only from the Nearctic Region where it is represented by the type species B. fidelis (Banks) and by B. speciosa (Banks). To date the biology of the immature stages of neither of these species has been elucidated. B. fidelis has been taken with fair frequency in the middle Atlantic states and the author has made several attempts to rear this interesting hemerobiid; however, it has only been recently that this effort has met with any success. Although this study represents a portion of a more general investigation of the morphology and ecology of the immature stages of the Nearctic Neuroptera, it seems desirable to publish these notes at this time because of their bearing on the ques- tion of the relationship of Boriomyia to the other genera of the Hemerobiidae. The writer wishes to express his gratitude to Mr. Ivan Huber of the Food and Drug Administration, Department of Health, Educa- tion and Welfare, who supplied the author with several living females, including the all-important specimens whose offspring form the basis for this account. * Methods The immature stages which were utilized in this study were de- rived from three females collected in College Park and Greenbelt, Prince Georges Co., Md., in 1956 and 1958. After rearing and pres- ervation, thirty-seven first instar larvae, ten second instar larvae, eighteen third instar larvae and two pupae were available for mor- phological investigations. Although this species can be identified from either sex of the adult with good reliability, the best taxonomic char- acters are found in the genitalia of the male. Accordingly, three pupae, the offspring of a single female, were allowed to transform and two of these produced males by which the initial identification of one of the original females was confirmed. These identifications were made by the writer utilizing the key of Carpenter (1940). 26 1960] MacLeod — B oriomyia 27 Past experience with numerous types of Neuroptera has indicated that when eggs can be obtained from field-collected females, these eggs are fertile. The usual procedure for inducing oviposition by such females has been to confine the insect in some suitable container. Thus Smith ( 1922) was quite successful in securing eggs from females of various Nearctic species of C hr y so pa by placing them in lamp- chimney cages over growing, aphid-infested plants, the aphids serving as food for the female and possibly also as an oviposition stimulus. Killington (1937) utilized much the same procedure in obtaining eggs from many species of British hemerobiids. Although this method was tried by the present writer, the inspec- tion of so large a volume for the infrequent eggs supplied by the females of B. fidelis proved difficult. In addition, the broadly oval wings of this species with their numerous macrotrichiae render the insect somewhat clumsy and almost any confined space serves to trap the female, the leaf axils of the plants serving especially well in this respect. A more satisfactory arrangement was found by confining the fe- males individually in 75 x 25 mm. cotton-stoppered glass vials. Hu- midity was supplied from a small pleget of cotton, dampened with a solution of sucrose and water which was placed in the bottom of the vial. To obviate the possibility of the female becoming stuck to the wet cotton, a cardboard partition was placed between the damp cot- ton and the female and this was removed only for a short time each day to permit the female to drink. Small nymphs of the aphid Mac- rosiphurn liriodendri (Monell) collected from nearby tulip trees ( Liriodendron tulipifera (L.)) were supplied on a small piece of leaf daily. Eggs, which were invariably laid on the cotton plug of the ovi- position vial, were removed as soon as detected and placed singly in 40 x 13 mm. cotton-stoppered shell vials. Larvae were supplied each day with nymphs of the same aphid as was fed to the adults. This food was introduced into the vial on a piece of fresh leaf which probably served to maintain a reasonable humidity although no at- tempt was made to control or record this variable. Small wads of damp cotton were placed in the vials after the cocoons had been spun in order to prevent excessive dryness; however, these were not placed in vials with larvae in order to eliminate any possibility of the larvae becoming trapped and drowning. These rearings were carried out in a room open to the usual fluctuations in the summer temperatures 28 Psyche [March - June which occurred during the periods of this study (June 25-August 8, 1956, and June 19-July 18, 1958). Color notes were made from microscopic observations of living larvae and these were supplemented by color photographs taken of freshly killed larvae. The drawings of Plates 5 and 6 were pre- pared by the author from these notes, from material fixed in Peter- son’s KAAD fluid (see Peterson, 1953, for details of preparation) and stored in alcohol, and from the examination of cleared specimens in temporary glycerine mounts with a binocular dissecting microscope and with a phase-contrast compound microscope. All measurements were made with an ocular micrometer calibrated with a stage mi- crometer. All indices of dispersion given in this paper are standard errors of the mean. Description of The Developmental Stages Oviposition and embryonic development — The three females used in this study were confined for periods of nine, fifteen and twenty-two days, during which times a total of ninety-four eggs was deposited. These were produced in a series of small lots which, with one excep- tion, contained between one and eight eggs (x = 3.53 ± 0.53 eggs/ lot). In a single instance a freshly caught female, whose abdomen was noted to be tremendously swollen with eggs, laid twenty-seven eggs during her first twenty-four hours of confinement. Subsequent to this, however, the egg production of this female fell within the limits noted above. In all cases oviposition occurred at night and was never observed by the writer. In addition to the ninety-four eggs produced by these females, the remains of an undetermined number of other eggs were noted which had, apparently, been partially consumed by the females. Six of the ninety-four eggs regarded as normal may also have been dam- aged by one of the females as these were slightly distorted in shape when first seen and none of them showed any signs of embryonic development. Three additional eggs although seemingly normal in all respects also failed to show any indication of development. Of the eighty-five eggs which showed apparent normal development, two failed to hatch. Egg — Approximately ellipsoid in shape, distinctly flattened on surface attached to substratum. Color, a pale greyish-yellow, becom- Explanation of Plate 5 Fig. 1. Facies of head capsule of first instar larva, dorsal view. Fig. 2. Head capsule and prothoracic “collar” of third instar larva, dorsal view. Psyche, 1960 Vol. 67, Plate 5 MacLeod Boriomyia 30 Psyche [March - June ing greyish-brown as embryo develops. Micropyle sessile, flattened, chalk-white; a minute depression in center. Chorion, apparently smooth, but showing a minutely granulate texture at a magnification of 6oX giving the impression of finely frosted glass. At higher mag- nifications an irregular imprinted pattern visible, attributed to the pressure of follicular cells of ovary by Killington (1946). Minute elevations and tubercles of chorion reported for several other hemer- obiid genera by Killington (1936) lacking in B. fidelis. Size of a single unhatched egg: length — 0.7752 mm., greatest width — 0.3420 mm. The eyes of the embryo become visible through the chorion after four days, while hatching takes place after about seven and one-half days of development (x = 7.6 zb 0.1 days, N = 70). As with other Neuroptera, emergence from the chorion is assisted by an egg- burster (figs. 5, 6) which forms a longitudinal ridge-like projection from the embryonic cuticle in the region of the clypeus. In a series of eggbursters derived from seven siblings which were studied in detail, the number of small teeth along the distal margin of the blade was found to vary between twenty-two and twenty-nine (x = 25.0 zb 0.9). The two extremes are figured in figs. 5 and 6. First instar larva (figs. 1, 3) — Head capsule ellipsoid, prominent, as wide as widest part of body; of a uniform grey color dorsally, this interrupted medially by a Y-shaped eedysial cleavage line passing down to clypeal margins just medial to bases of jaws on either side; anterior, diverging arms of cleavage line cross connected by trans- verse area of pale cuticula at level of anterior tentorial pits delimit- ing a proximal frontal area and a distal clypeal area. Ocular area of head capsule antero-lateral, consisting of three, clear corneal swell- ings. Anterior two corneae each with one dark ommitidial element beneath ; three additional ommatidiae visible beneath posterior cornea. Latero-ventral surface of head capsule with usual pair of dark fuscous lines; the more dorsal, running between posterior tentorial pit and dorsal articulation of mandible, representing the subgenal sulcus, the more ventral, running from vicinity of posterior tentorial pit to maxil- lary component of jaw, representing cardo and stipes. Ventral surface of head capsule largely covered with shield-shaped mental region of labium. Antenna three segmented, short, reaching only to tip of extended jaws. Basal segment dark, fuscous; short, only slightly longer than wide. Second segment comprising most of length of antenna, about five times as long as first segment. Third segment short, peg-like, 1960] MacLeod — Boriomyia 3i approximately of same length as basal segments; surmounted by a terminal seta slightly longer than segment itself. Distal two segments pale grey, showing usual irregular pattern of annular sclerotizations characteristic of antennae of hemerobiid larvae (fig. 1). Jaws slightly curved inwards; pale grey toward base, becoming dark brown for distal one-third of length. Labial palpi three segmented. Basal seg- ment short, slightly wider than long; arising from largely fuscous, mound-like palpigeral swelling on antero-ventral surface of head capsule. Second segment of same shape as first segment. Both of basal two segments braced by a mid-segmental thin, dark, sclerotic ring. Distal segment of palpus elongate, inflated, reaching well be- yond tips of jaws when extended forwards. Surface of distal palpal segment covered with same irregular pattern of annular sclerotiza- tions found on distal two segments of antenna. Base of distal palpal segment braced by a thin fuscous ring. Thorax — Prothorax with usual three subdivisions. Anterior subsegment short, transverse, forming a collar which overlaps posteri- or margin of head capsule only slightly. Posterior subsegment of prothorax with spiracle opening on lateral surface. Mesothorax with two subdivisions, metathorax consisting of leg-bearing portion only. Leg-bearing portions of thoracic segments with latero-dorsal sclerites reduced to mere vestiges, easily visible only upon examination of cleared specimens with phase microscope. Legs largely pale except for thin fuscous sclerotizations partially encircling distal end of each coxa and each femur and proximal mar- gin of each tibia. Pretarsus of all legs with usual clavate empodium extending from between the pretarsal claws. Abdomen consisting of ten segments; first eight segments mem- branous, with a spiracle located laterally on each segment. Segment nine with a sclerite covering most of dorsal surface. Tenth abdominal segment largely covered with four small sclerites, one dorsally, one ventrally and one on each side. A pair of eversable pygopods con- tained within posterior portion of this segment. Cuticle of thoracic and first eight abdominal segments of unfed larva a light pinkish-purple color through which color of body con- tents shows through as a light greenish-yellow background. Color of this background changing with the intake of food to an orange hue and then to a dark reddish-orange extending from middle of prothorax to second abdominal segment. Abdominal segments nine and ten pale grey throughout the first larval stadium with their sclerites just visible under a dissecting microscope. Aorta at first only 32 Psyche [March - June faintly visible between anterior prothorax and eighth abdominal seg- ment, toward end of stadium becoming outlined by small isolated patches of developing fat body on either side of aorta. Fat body particularly well-developed in posterior subsegments of pro- and mesothorax. Duration of first larval stadium — 3.4 zb 0.7 days (N = 33) Length of unfed first instar larva (measured from clypeal margin of head capsule to posterior margin of dorsal sclerite of tenth abdominal segment) — 1.57 zb 0.12 mm. (N = 6) Maximum width of head capsule (measured across ocular areas) — 0.3018 zb 0.0009 mm. (N = 7) Subsequent development of B. fidelis — The head capsules of the second and third instar larvae (fig. 2) are quite similar to each other and differ from that of the first instar larva principally in being slightly wider in proportion to their lengths, which, combined with a lack of the curvature of the lateral margins of the head present in the first instar larva, gives the dorsal aspect of the head capsule a distinct- ly more quadrate appearance. In addition, the formerly fuscous regions of the clypeal area dorsally and the mental region of the labium ven- trally of the first instar larva are pale in the subsequent larval instars so that only the frontal and parietal areas are darkened in these stages. The labial palpi are somewhat less inflated in appearance in the sec- ond and third instar larvae although they remain stouter than is usual in this family and reach well beyond the tips of the extended jaws. The pretarsi of all legs of the second and third instar larvae lack the empodium present in the first instar larva, its place being taken by a delicate, fan-like arolium, arising from the dorsal surface of the pretarsus. Posteriorly, the tergal area of the nineth abdominal seg- ment is no longer covered by a distinct sclerite in the second and third instar larva. The body form of the second and particularly the third instar larva (fig. 4) becomes progressively swollen and grub-like with the develop- ment of a complex series of projecting lobes on the original segments of the body. The anterior subsegment of the prothorax remains short and collar-like and the head capsule becomes increasingly retracted within this portion of the body, so that although the head can be ex- tended, the usual condition in which to find a living larva is with the 1960] MacLeod — Boriomyia 33 head withdrawn within the prothorax as far anteriorly as the bases of the antennae. The body pattern visible to the eye is the result of the interplay between three somewhat variable features. The integument is lightly pigmented by a mottled pattern of a diffuse, purple color which over- lies a much darker reddish-purple color due to the gut contents and hemolymph. Between these regions, the rapidly developing fat body creates an intricate series of patches which are nearly pure white in regions where the cuticle is faintly pigmented and which show as darker purple areas where the cuticle is more darkly pigmented. The exact pattern which the larva presents depends largely on the extent of the development of the fat body which gradually fills most of the available space of the hemocoel. One constant feature of this pattern is the presence of two white transverse bands across the body on the posterior subsegments of the prothorax and mesothorax due to the near lack of cuticular pigment in these regions. Ventrally the pal- pigers consistently include a small portion of the fat body within them which shows through the clear cuticle as a snow-white patch. In a dorsal view of a fully mature third instar larva (fig. 4) the fat body will be seen to have ramified throughout the body leaving only paired lateral “windows”, from the posterior margin of the prothorax through the sixth abdominal segment, and the aorta in the dorsal mid- line through which the darker color of the hemolymph shows through. A typically hemerobiid cocoon is spun by the third instar larva. This consists of a loose network of silk strands within which is located a more-compact inner cocoon which is, nevertheless, open enough to permit easy observation of the final stages of development. Pupation, metamorphosis and the eclosion of the adult all occur with- out any apparent cessation of development. The pupa is typical of that of many hemerobiids studied by the writer and, so far as can be ascertained, possesses no prominent features peculiar to Boriomyia. Post first instar development may be summarized as follows: Measurements Length mature second instar larva — mature third instar larva — Maximum width of head capsule second instar larva — third instar larva — Developmental times second instar larva — 3.32 zb 0.43 mm. (N = 5) 4-53 ± 1-05 mm. (N = 5) 0.3426 zb 0.0063 mm. (N = 4) 0.4073 zb 0.0087 mm. (N = 3) 3.5 zb 0.7 days (N = 20) 34 Psyche [March - June third instar larva — prior to spinning cocoon — 4.5 ± O.i days (N = 4) post spinning — 6.9 zb 0.4 days (N = 4) pupa — 10.7 zb 0.4 days (N = 3) Behavioral and ecological notes — The young first instar larva is the only stage which is really active in the sense of seeking out prey. The older first instar larva and particularly the second and third instar larvae tend to remain immobile in some corner of the vial, a favored site for this position being between the cotton plug and the side of the vial. Aphids which move too close to the larvae are seized by a quick upward thrust of the jaws from below. The prey does not seem to be sucked as thoroughly as has been observed to occur with various species of Hemerobius and Micromus with which the author has worked. In these genera the prey is manipulated by the larva until the jaws have probed into every available corner of the body. The larvae of B. fidelis seem to simply suck the immediately available fluids of the prey with only incidental movement of the aphid. One behavioral trait of the adult deserves some comment because of its bearing on the question of the affinities of Boriomyia discussed below. When at rest, the adult consistently assumes a distinctive position by placing the anterior surface of the head flat against the substratum with the antennae directed posteriorly between the legs and neatly clasped by both pairs of palpi. In many respects this position is reminiscent of the death-feigning position which many Neuroptera will assume if startled suddenly by a sharp vibration, although in this case the insect drops to the ground and lies on its side and even though the antennae are directed posteriorly beneath the body, they are not clasped by the palpi. The natural ecology of this insect remains almost completely un- known as the writer has yet to locate any of the immature stages in the field. Of possible interest in terms of the habitat of this species is a collection of three males and two females taken in a short time in an open pine woods near Charleston, S. C. These were captured on April 20, 1957, and I am informed by the collectors. Messers. Ivan Explanation of Plate 6 Fig. 3. First instar larva, dorsal view. Fig. 4. Mature third instar larva, dorsal view. The latero-dorsal sclerites of the larvae shown in figs. 4 and 5 are depicted as seen by the microscopic examination of cleared specimens. In life these sclerites are nearly invisible. Figs. 5 and 6. Egg bursters, showing extremes in the number of teeth, lateral view. Psyche, 1960 Vol. 67, Pi. ate 6 MacLeod — Boriomyia 36 Psyche [March - June Huber and Paul E. Spiegler, that the insects seemed to be associated with piles of dead branches of yellow pine ( Pinus australis Michx.) on the ground. The Taxonomic Status of Boriomyia A series of nomenclatorial mistakes combined with several taxono- mic errors in evaluating published descriptions have obscured the true relationships of this genus within the Hemerobiidae. The name Boriomyia appeared for the first time in 1904 when Banks included his previously described species PTemerobius fidelis and H. speciosus under this generic name. A formal description of Boriomyia as a new taxon appeared in 1905 in Banks’ Revision of the Nearctic Hem- erobiidae. In this work, six Nearctic species were included in this genus in addition to B. fidelis and B. speciosa, one of these, H. dis- junctus Banks, being designated as the type species. Subsequently additional Nearctic and numerous exotic species were referred to Boriomyia by various workers. In 1930 Banks, recognizing that B. fidelis and B. speciosa did not form a homogeneous group with the remainder of the species placed in Boriomyia, erected a new subgenus, Allotomyia, for these two species and subsequently (1935) treated this group as a full genus. Killington (1937) agreed that the two groups of species recognized by Banks should be separated generically, but pointed out that since the name Boriomyia had been validated in 1904 when it was used in combination with two already-described species, Banks’ 1905 designa- tion of H. disjunct us as the type of Boriomyia was in violation of the Rules since this species was not included in Boriomyia when this name was originally validated. Designating H. fidelis as the type species of Boriomyia, Killington proposed the name Kimminsia for the British species formerly included in Boriomyia, the remaining non-British species of the former Boriomyia (other than B. speciosa) presumably being also referable to this genus. A Palaearctic species, H. betulina Str0m was selected as the type of Kimminsia. These two genera were considered to be separated by venational features which had already been noted by Banks. These views were reiterated by Killington in 1937b. Tjeder (1941) in a critique of Killington’s work and with- out specimens of B. fidelis to study felt that the venational distinctions which had been drawn between Boriomyia and Kimminsia were not sufficient to warrant generic separation. Subsequently in a long series of papers ( 1943a, 1943b, 1944, 1945, I95E 1953a, 1953b, 1954, 1 955 , i960) , he has continued to use the name Boriomyia to cover the Kimminsia group of species. 1960] MacLeod — Boriomyia 37 Circumstances have thus conspired to make it seem that Kirnminsia is a taxonomic segregate from the closely allied genus Boriomyia and that the question of whether to recognize one genus or two simply depends on the emphasis which one places on the differences in the venation of these two groups. Actually this is misleading, for as long ago as 1940 Carpenter pointed out that the differences between these two genera are much more far-reaching than had been suspected, in- volving not only consistent differences in venation, but fundamental differences in the basic organization of the genitalia of both sexes. Indeed, it was Carpenter’s contention that the true affinities of Bori- omyia lie not with Kirnminsia at all, but rather with the genus Me- galomus. The findings of the present study lend complete support to this view. Details of the biology of the Palaearctic Megalomus hirtus (Linne) including several excellent figures of the larvae have been published by Killington (1934, 1937a), while the present author has reared the immature stages of an unidentified species of Megalomus from Mexico. In addition, modern figures and descriptions of larvae of species of Kirnminsia, W esmaelius, Psectra, Sympherobius, Hetne- robius, Micromus, and Drcpanepteryx have also been provided by Smith (1923, 1934), Killington (1936, 1937a, 1946) and Fulmek (1941) and as the writer has been able to rear or study larvae of species of the last four of these genera, detailed comparisons of a variety of larval types of the Hemerobiidae have been possible. Only in very general features shared by the larvae of all known hemerobiids do those larvae of Micromus, Hemerobius , W esmaelius and Kirnminsia which are known bear any close structural resemblance to the larvae of the remaining groups and they will not be discussed further. A series of striking similarities is to be found in the larvae of Boriomyia and Megalomus involving the shape of the labial palpi, the form of the terminal segment of the antenna and in the appearance of the mature larva. The swollen, inflated shape of the labial palpi, particularly striking in the first instar larvae, but noticeable in the later two instars as well, is present only in Sympherobius and Psectra in addition to Megalomus and Boriomyia although it has never been observed to reach the extreme degree in the first two of these genera that it does in the latter two. The extremely shortened terminal seg- ment of the antenna, surmounted by an apical seta nearly as long as or longer than this segment, is known to the writer only in Boriomyia and Megalomus. A much less drastic reduction in the terminal seg- ment occurs in Sympherobius, but here the segment is somewhat flask- 38 Psyche [March - June shaped as it is dilated basally and the terminal seta is much shorter than the length of the segment. The swollen, physogastric appearance of the mature larva has probably been produced several times inde- pendently in the Hemerobiidae in correlation with a more sessile mode of larval life. The condition has been approached in Sympherobius , Psectra and Drepanepteryx although in no species of these groups known to the writer does it reach the extreme which has been pro- duced in Boriomyia and Megalomus. Associated with this inflated, sessile larva is the habit of carrying the head deeply retracted within the prothorax, developed to an extreme degree in Boriomyia, Mega- lomus, and Drepaneptaryx, and the supression of the latero-dorsal sclerites of the thoracic segments. In Sympherobius, Psectra and Drepanepteryx these sclerites have been strongly reduced on the meso- and metathorax, but those of the prothorax have remained relatively large and prominent. In Boriomyia and Megalomus alone have the prothoracic latero-dorsal sclerites been reduced to the point where they are difficult to demonstrate. Killington states that these sclerites are lacking on the prothorax of M. hirtus although it seems possible that close scrutiny might reveal their presence since their remnants can just be detected in B. fidelis as well as the Mexican species of Megalomus studied by the writer. One additional point of resemblance between members of Boriomyia and Megalomus is to be found in the peculiar rest position of the adult noted above for B. fidelis, which was observed by the writer in adults of the Mexican species of Megalomus studied, and noted by Killington (1937a) in M. hirtus. The writer has presented the evidence for the relationship of Bori- omyia and Megalomus to Mr. Tjeder who, it has been noted, pre- viously felt that there were insufficient characters to separate Kim- minsia from Boriomyia. In addition, specimens of B. fidelis were supplied for his study. Mr. Tjeder has informed the writer (in lift.) that he is now in full agreement with the placement of B. fidelis near Megalomus (indeed, he feels that this species must be placed in Megalomus) . He has, however, continued to use the name Boriomyia for the Kimminsia species (Tjeder, i960), arguing (in litt.) that Banks’ genotype designation for Boriomyia in 1905 must be considered valid. The writer is of the opinion that the Rules are perfectly clear on this point and that the validation of Banks’ designation would re- quire an action of the International Commission to set aside the Rules. It should be noted that in addition to Killington (1937a, 1937b) and Carpenter (1940), Fraser (1940, 1942, T959), Friedrich ( 1 953 ) > 1960] AlacLcod — Boriomyia 39 Nakahara (1956, i960) and Parfm (1956) have used the names Boriomyia and Kimminsia in the sense adopted in the present paper. Concerning the question of the validity of Boriomyia as a genus distinct from Megalomus, this must await a study of the type species of Megalomus , the Palaearctic M. tortricoides Rambur, as well as a wider variety of species of this genus than has been available to the present writer. Carpenter (1940) was able to separate these genera on the basis of the position of the radial cross vein with reference to the point of origin of R5 in the hindwing, although he felt that species which were intermediate in this character would probably be found. Recently Nakahara (i960) has observed that B. fidelis possesses a median lobe of the fused parameres of the male genitalia (present also in B. speciosa) which is said to be lacking in species of Megalomus. A second character of B. fidelis considered by Nakahara to be of gen- eric importance, the basally fused and apically bifucrate “aedeagus”, is present also in at least the Nearctic species M. minor Banks which is a Megalomus by the criterion of a lack of the median lobe of the fused parameres. It is the opinion of the present writer that stable generic limits in the Hemerobiidae usually involve greater differences in the basic ground plan of the male genitalia than those aduced by Nakahara, and frequently correlate with differences in the structure of one or more of the immature stages and in constant venational features as well. In these respects, Boriomyia seems poorly distinct from Megalo- mus and will probably fall as a synonym of this genus. Literature Cited Banks, N. 1904. A list of the neuropteroid insects, exclusive of Odonata, from the vicinity of Washington, D.C. Proc. Ent. Soc. Wash. 6:201-217. 1905. A revision of the Nearctic Hemerobiidae. Trans. Amer. Ent. Soc. 32:21-51. 1930. New neuropteroid insects from the United States. Psyche 37:223-233. 1935. A few new North American Neuroptera. Psyche 42:53-57. Carpenter, F. 1940. A revision of the Nearctic Hemerobiidae, Berothidae, Sisyridae, Polystoechotidae and Dilaridae (Neuroptera). Proc. American Acad. Arts Sci. 74:193-280. Fraser, F. 1940. Kimminsia rava (Withy.) in Hampshire. Entomologist 73: 166. 1942. A new species of Kimminsia (Neur., Hemerobiidae), with historical figures and notes. Ent. Monthly Mag. 78:80-86. 1959. Mecoptera, Megaloptera and Neuroptera. Handbooks for the Identification of British Insects 1(12-13) :l-40. London: Royal Ent. Soc. London. Friedrich, H. 1953. Neuroptera. Bronns Klassen und Ordnungen des Tier- reichs. Funfter Band, 3. Abteilung, XII. Buch, Teil a. Leipzig: Akademische Verlagsgesellschaft Geest & Portig K.-G. pp. 1-148. 40 Psyche [March - June Fulmek, L. 1941. Uber die Aufzucht von Drepanepteryx phalaenoides L. ex ovo. (Neuroptera: Planipennia, Hemerobiidae) . Arbeiten uber Morphology und taxonomische Entomologie aus Berlin-Dahlem 8:127-130. Killington, F. 1934. On the life-histories of some British Hemerobiidae. Trans. Soc. British Ent. 1:119-134. 1936. A monograph of the British Neuroptera. Volume I. London: The Ray Society, pp. 1-269 + i-xix. 1937a. A monograph of the British Neuroptera. Volume II. London: The Ray Society, pp. 1-306 + i-xii. 1937b. The generic names of the British Neuroptera, with a check list of the British species. The Generic Names of British Insects 4:60-80. London: Royal Ent. Soc. London. 1946. On Psectra diptera (Burm.) (Neur., Hemerobiidae), including an account of its life history. Ent. Monthly Mag. 82:161-176. Nakahara, W. 1956, New or little known Neuroptera from Japan and adjacent Territories (Neuroptera). Kontyu 24:182-191. 1960. Systematic studies on the Hemerobiidae (Neuroptera). Mushi 34:1-69. Parfin, S. 1956. Taxonomic notes on Kimminsia (Neuroptera: Hemero- biidae). Proc. Ent. Soc. Wash. 58:203-209. Peterson, A. 1953. A manual of entomological techniques. Ann Arbor, Michigan: J. W. Edwards, pp. 1-367 + i-v. Smith, R. 1922. The biology of the Chrysopidae. Cornell Univ. Agric. Expt. Sta. Memoir 58:1287-1372. 1923. The life histories and stages of some hemerobiids and allied species. Ann. Ent. Soc. Amer. 16:129-148. 1934. Notes on the Neuroptera and Mecoptera of Kansas, with keys for the identification of species. Jour. Kansas Ent. Soc. 7:120-144. I'jeder, B. 1941. Some remarks on “The generic names of the British Neu- roptera’’. Ent. Tid. 62:24-31. 1943a. The Neuroptera and Mecoptera of Northern Norway (Nordland, Troms and Finnmark). Troms Museums Arshefter, Naturhis- torisk Avd. Nr. 25. Vol. 63 (1940) : 1 - 1 5. 1943b. Revision of the Norwegian Neuroptera and Mecoptera. Norsk Ent. Tid. 6:133-139. 1944. Norwegian Neuroptera and Mecoptera in the Bergen Mu- seum. Bergens Museums Arbok, Naturvitenskapelig rekke Nr. 1:1-11. 1945. Catalogus Neuropterorum et Mecopterorum Norvegiae. Norsk Ent. Tid. 7:93-98. 1951. The mountain fauna of the Virihaure area in Swedish Lapland. Neuroptera. K. Fysiogr. Sallsk. Lund Handl. N. F. 61:121. 1953a. Faunistical notes on Swedish Neuroptera. Opus. Ent. 18: 227-228. 1953b. Catalogus insectorum Sueciae. Additamenta ad partes I-X. Opus. Ent. 18:71. 1954. Genital structures and terminology in the order Neurop- tera. Ent. Medd. 27:23-40. 1955. Two new species of Boriomyia from South Africa. Annals South African Mus. 51:381-385. 1960. Neuroptera from Newfoundland, Miquelon, and Labrador. Opus. Ent. 25:146-149. THE FEMALE OF IN OCELLI A PILICORNIS CARP. (NEUROPTERA: INOCELLIIDAE) P By F. M. Carpenter Harvard University The snakefly, Inocellia pilicornis Carpenter, was described in 1959 (Psyche, 65: 56-58) from three males collected in the states of Nuevo Leon and Tamaulipas, Mexico. These males were unique among known Raphidiodea in possessing moniliform antennae, their seg- ments bearing long radiating setae, arranged in a definite pattern. In July, i960, Dr. H. F. Howden, who collected one of the males of the original set, found the first known female of this species, along with another male, very near the type locality. It now turns out that the female also has moniliform antennae, but the hair covering of the antennae, although distinctly different from that of other species, is not as similar to that of the male of pilicornis as might be expected. The color markings of the female are like those of the male type, except that the median white marks on the meso- and metanotum and the abdominal tergites are in the form of patches instead of a contin- uous stripe. The forewing is 6.8 mm. long and 1.8 mm. wide; the Figure 1. Portions of antennae of Inocellia pilicornis Carp, a, female, x 140 (specimen from Chipinguy Mesa) ; b, male, x 55 (holotype). pterostigma, having a maximum length of 1 mm. and a width of a little less than .5 mm., is relatively shorter than in the male. The wing venation is like that of the holotype, illustrated in my account 1 Published with the aid of a grant from the Museum of Comparative Zoology at Harvard College. 41 42 Psyche [March - June of the species. The antennae of the female are 3.7 mm. long. Each consists of 42 segments, instead of 60 as in the male, and the individu- al segments are smaller than in the male. The hairs on the antennal segments are shorter than those of the male and form a different pat- tern (figure 1). A ring of twelve conspicuous setae is located near the distal end of each segment and two prominent hairs occur on one side of each segment, slightly distal of the ring just mentioned; these two hairs are a little longer than the hairs comprising the ring and they almost always project in the same direction, as shown in figure 1 a. The more proximal part of each antennal segment includes about 20 short hairs which are arranged somewhat irregularly and which do not form a second ring, as in the male. The ovipositor of the female is like that of other Inocellia ; in the specimen at hand it is 5.5 mm. long. The body length of the female, exclusive of the anten- nae and ovipositor, is 6 mm. It seems virtually certain that this new female, which was collected at Chipinguy Mesa, near Monterrey, Nuevo Leon, Mexico, July 28, i960, belongs to pilicornis. A male, collected by Dr. Howden at the same time, is like the holotype, except that the white markings on the abdominal tergites are somewhat smaller. Although the female turns out to have antennal characteristics as distinctive as those of the male, I do not believe that a new genus is necessary for this species. Other features, including the abdominal structures of the male and female, are similar to those of other species of Inocellia . A NOTE ON THE PREY AND A NESTING SITE OF CERCERIS TRUNCATA CAMERON (HYMENOPTERA: SPHECIDAE: PHILANTHINAE) By F. G. Werner Department of Entomology, University of Arizona This species seems to be very rare or extremely local in southern Arizona; Dr. H. A. Scullen, who has kindly provided the identifica- tion, held it as undescribed until he discovered that it had been described from Mexico by Cameron. However, there is a rather extensive nesting site in my own yard, at 1247 N. Warren Ave., Tucson. During the past four years I have been able to observe the habits of the species regularly and have found that the females store only beetles of the family Bruchidae ( Mylabridae) . The species stored at this site are all of medium size, and probably all come from leguminous trees and shrubs in the neighborhood. In order of abun- dance the bruchids gathered are: Algarobius prosopis (Lee.), Ali/no- sestes protractus (Horn) , Mimosestes amicus (Horn) and N eltumius arizonensis (Schffr.) ; this is roughly the relative abundance of these species when they are taken in general collecting. The most abundant source tree in the neighborhood is the so-called Mexican palo verde, Parkinsonia, which is planted extensively, blooms in the spring and has mature pods by mid-July, when the wasps become active. The site occupied is on fiat ground in the southwestern corner of the lot, about 10' x 10', shielded on south and west by a five-foot wall and partly shaded by large oleander bushes ; the soil is bare and con- sists of well-packed fill about eight inches deep over caliche ; almost all of the water that reaches it comes from rainfall. Activity of the wasps was checked weekly in 1958, when the first swarms of males and starts of nests were observed on July 20 and could not have started more than a week before this date. The males fiy continually during the day, just over the oleander bushes and nearby vegetation, stopping occasionally to rest. Hundreds of individuals are involved in these flights. Smaller numbers, usually about twenty, patrol the nesting site, flying about four inches above the ground, and attempt copulation with females flying in the area or returning to their nests with prey. I could not discover where the males went at night. The females dig small holes, either bare or with a low mound up to ij/2" in diameter. Most of their flying and provisioning activity occurs in the morning but even then the principal activity seems to consist of sitting in the nest entrance with only the front of the head exposed. By noon all 43 44 Psyche [March - June is quiet and most of the nest entrances are loosely plugged, only the occasional pushing up of dirt in some of the nests indicating that dig- ging may be in progress. Marked nests were open for two or three days, except when they were plugged with loose dirt in the afternoon, and one nest was open at least five days. The nests were hard to keep track of during the period of observation because of frequent rains. By July 27 there were 55 nests in a marked area 4' x 4' and there were at least this many active during the rest of the summer. Solenopsis ants raided part of the site but there was no sign of social or other parasites. Individual nests were dug up but none could be excavated com- pletely. The holes could be followed down to about 4", where they seemed to end in loose dirt ; below some of them, at 5" to 6", a cluster of up to ten bruchids, with an egg or larva, indicated where the cell must have been. The cells must be of very loose construction. Nesting activity and male swarming continued from mid- July well into Sep- tember in 1958, gradually diminishing during the latter month. A similar schedule has been maintained in the two succeeding years. No adults have been seen until July, the normal starting time of the summer monsoon season. THE TYPE SPECIES OF THE ANT GENUS EURHO- PALOTIIRIX . — In our paper, “A world revision of the ant tribe Basicerotini,” Studia Entomologica, Petropolis, Brazil, 3: 202, i960, we erected a new genus Eurhopalothrix to receive certain species formerly placed in Rhopalothrix Mayr, along with some new species. Through inadvertance, no type species was cited for the new genus, and we therefore here designate Eurhopalothrix bolaui (—Rhopalo- thrix holaui Mayr, 1870) as type species of the genus Eurhopalothrix Brown and Kempf. — W. L. Brown, Jr., Department of Entomol- ogy, Cornell University, Ithaca, N. Y., and W. W. Kempf, o. f. m., Sao Paulo, Brazil. CAMBRIDGE ENTOMOLOGICAL CLUB A regular meeting of the Club is held on the second Tuesday of each month October through May at 8:00 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 shows the head of the worker of Myrmoteras karnyi Gregg. The original, drawn by Dr. R. E. Gregg, was published in Psyche, Volume 61, p. 21, 1954. BACK VOLUMES OF PSYCHE The Cambridge Entomological Club is able to offer for sale the following volumes of Psyche. Volumes 3, 4, 5, 6, 7, 8, each covering a period of three years. $8.00 each. Volumes 10, 14, 17 to 26, each covering a single year, $2.00 each. Volumes 27 to 53, each covering a single year, $2.50. Volumes 54 to 65, each covering a single year, $3.00. Volume 66, covering a single year, $5.00. Some other volumes, lacking certain issues, are also available (information upon request). Orders for 2 or more volumes subject to a discount of 10%. Orders for 10 or more volumes subject to a discount of 20%. All orders should be addressed to F. M. Carpenter, Editor of Psyche, Biological Laboratories, Harvard University, Cambridge, Mass. 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 38. Mass. PSYCHE A JOURNAL OF ENTOMOLOGY Established in 1874 Vol. 67 September, i960 No. 3 CONTENTS A Study of Bembix u-scripta, a Crepuscular Digger Wasp. H. E„ Evans 45 Defense Mechanisms of Arthropods. II. The Chemical and Mechanical Weapons of an Earwig. T. Eisner 62 A Triassic Odonate from Argentina. F. M. Carpenter 71 On the True Nature of the Azygethidae (Chilopoda: Geophilomorpha) . R. E. Crabill, jr. 76 CAMBRIDGE ENTOMOLOGICAL CLUB Officers for 1960-61 President H. Levi, Harvard University Vice-President N. S. Bailey, Bradford Junior College Secretary G. L. Bush, Harvard University Treasurer F. M. Carpenter, Harvard University Executive Committee A. G. Humes, Boston University J. J. T. Evans, Harvard University EDITORIAL BOARD OF PSYCHE E. M. Carpenter (Editor), Professor of Entomology , Harvard University P. J. Darlington, Jr., Head Curator of Insects, Museum of Com- parative Zoology W. L. Brown, Jr., Assistant Professor of Entomology , Cornell University ; Associate in Entomology, Museum of Comparative Zoology E. 0. Wilson, Associate Professor of Zoology, Harvard University H. Levi, Associate Curator of Arachnology , Museum of Comparative Zoology 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, $5.00 to all other subscribers. Single copies, $1.25. Checks and remittances should be addressed to Treasurer, Cambridge Ento- mological Club, Biological Laboratories, Harvard University, Cambridge, Mass. Orders for back volumes, missing numbers, notices of change of address, etc., should be sent to the Editorial Office of Psyche, Biological Laboratories, Har- vard University, Cambridge, Mass. IMPORTANT NOTICE TO CONTRIBUTORS Manuscripts intended for publication should be addressed to Professor F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. Authors contributing articles over 8 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 $6.00 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, $18.00 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 i960 Psyche (Vol. 67, nos. 1-2) was mailed Feb- ruary 16, 1961. The present issue of Psyche (Volume 67, no. 3) is published with the aid of a National Science Foundation Grant (G 15817) to the Cambridge Entomological Club. The Lexincton Press. Inc., Lexington, Massachusetts Vol. 67 No. 3 PSYCHE September, i960 A STUDY OF BEMBIX U-SCRIPTA , A CREPUSCULAR DIGGER WASP By Howard E. Evans Museum of Comparative Zoology One of the most characteristic features of the digger wasp genus Bembix is the almost complete obliteration of the ocelli. Two North American species, arcuata and u-scripta, provide the only important exceptions to this, and these species also possess several other structural features which set them aside from all other species of Bembix. A few years ago I presented some very incomplete data on a colony of u-scripta in southern Texas, suggesting at the same time that the be- havior of the species ought to prove of unusual interest because of its apparently primitive structural features (Evans 1957)- I have recently made further studies of this same colony and also of another small colony in southwestern New Mexico, thus providing an unusual opportunity to compare the behavior of two widely separate colonies of an uncommon species of distinctly localized distribution. 1 hese studies also demonstrate that u-scripta has one very remarkable feature in its behavior: the females hunt and provision only in the dusk, around and after sunset. Throughout the higher Hymenoptera, large ocelli are distinctly correlated with nocturnal flight. It seems very probable that the preservation of functional ocelli in this species repre- sents an adaptation for flying in the twilight. A second problem concerns the identity of Bembix arcuata : is this in fact a distinct species from u-scripta ? The Texas colony was located not too far from the type locality of arcuata, the New Mexico colony not far from that of u-scripta. The New Mexico wasps had a much more extensively yellow color pattern (more like u-scripta) , but there was considerable color variation in wasps from both localities. In fact, on the basis of the Texas locality alone, I have already tenta- tively placed arcuata in the synonymy of u-scripta (Evans 1959)- Clearly it will be of interest to see what light behavior sheds on this matter. In this paper I shall first consider the structural peculiarities of Bembix u-scripta and also the differences which are supposed to sepa- 45 ^THSONIAN ■^BTHTOTION 46 Psyche [September rate u-scripta and arcuata, since this information lends more significane to the behavioral studies. There follows a discussion of the ethology of the two colonies studied and finally a brief statement of conclusions. T he studies of the Texas colony were made with the assistance of a grant from the National Science Foundation, while those of the New Mexico colony were made during tenure of a fellowship from the John Simon Guggenheim Memorial Foundation. I am indebted to Dr. M. A. Cazier, Director of the Southwestern Research Station of the American Museum of Natural History, for making available the facilities of that station during the summer of 1959. I am also indebted to Drs. Cheng Shan Lin and Oliver S. Flint for as- sisting me in the field work on the Texas colony. The dipterous prey of the Bembix have been identified by the following specialists, to all of whom I am much indebted: W. L. Downes, R. H. Foote, M. T. James, C. Martin, R. H. Painter, L. L. Pechuman, H. J. Reinhard, C. W. Sabrosky, Y. Sedman, H. V. Weems, and W. W. Wirth. Morphological Considerations The important structural features of Bembix u-scripta and arcuata are given as follows by Parker (1917, 1929) : 1. Ocelli unusually well developed and apparently functional (posterior ocelli with lenses clear, convex, circular except truncated laterally; anterior ocellus in form of a fairly wide transverse band). 2. Second transverse cubital vein of front wing only slightly curved (in other species of Bembix it is sharply bent). 3. Male with basal segment of middle tarsus distinctly curved, its inner surface beset with several stout spines; middle femora of male dentate, middle tibiae ending in a spine-like process. 4. Male with abdominal sternites devoid of processes, but seventh with a pair of carinae which diverge basally. 5. Labrum with a small median elevation (both sexes). 6. Basal segment of front tarsus with an unusually large number of comb-spines (8-10) (both sexes). 7. Scape of male much thickened. 8. Pygidium of female slightly rugose and with strong lateral ridges. 9. Pattern of yellow maculations unusually strongly developed. To these nine characters should be added another: 10. Apical tergite of male with deep longitudinal lateral incisions which cut off a pair of appendages which much resemble the parameres of the genitalia. 1960] Evans — Bembix u-scripta 47 All of these features are unique or shared with only a very few other species of Bembix. Some of them, though unique in Bembix, are shared (at least in some measure) with species of the more primi- tive bembicine genus Stictiella ; these include characters I, 2, and 3 above. Lohrmann (1948) has studied coloration in Bembix at some length, and has concluded that reduction in color pattern (i.e., less yellow) is correlated with structural advance; thus, on these grounds, character 9 should also be regarded as primitive. The remaining six characters should be regarded as specializations within the genus. However, those which involve the female (5, 6, 8) are not absolutely unique and the remainder (4, 7, 10) involve male secondary sexual characters, which are remarkably plastic in the bembicine wasps. 1 he more striking features thus suggest that these wasps do, in fact, retain several features which may have been characteristic of the ancestral Bembix , though obviously an assortment of specializations of a less striking sort have been acquired. In describing Bembix arcuata, Parker (1917) listed the following characters by which it differed from u-scripta (Fox 1895) : 1. The nature of the male “genital stipes” (i.e., parameres), which are much more slender and attenuate than in •. u-scripta . 2. Mesosternum marked with black (yellow in u-scripta) . 3. Abdominal sternites 2-4 with small lateral yellow spots (in u-scripta these are more extensively yellow and there is some yellow on the sternites behind these). 4. Apical sternite black (yellow in u-scripta ). Since the first of these characters involves an important structural distinction, it should be considered first. Parker based his interpreta- tion of u-scripta on the type specimen, one other male, and four fe- males. I have examined these two males and find that it is true that the parameres are less attenuate than in the type of arcuata — or in males from both colonies studied. However, the difference is slight and relative, and I cannot agree with Parker that the genitalia “differ widely”. Parker’s figures tend to exaggerate the differences between the two: yet the differences cannot be entirely reasoned away. Study of the color characters supposed to separate these species re- veals that while they may apply to the type specimens, they are by no means absolute when additional specimens are considered. The type of u-scripta is almost wholly yellow, that of arcuata predominantly black. Specimens from the Texas colony which I studied are close to arcuata , but exhibit enough variation to cause confusion in using Parker’s key. 48 Psyche [September Specimens from the New Mexico colony are close to u-scripta, but again they do not agree perfectly and there is much variation to be considered. When specimens from other localities are considered also, it becomes obvious that the extent of maculation varies in a complete spectrum all the way from “arcuate.” to “u-scripta” . I have tabulated the variation in two color characters (Tables I and II), selecting these two merely because Parker emphasized them. I have also studied other characters, such as the very noticeable variation in the develop- ment of the U-shaped markings on the mesoscutum, with very similar results. This species has not been collected in enough localities to en- able one to be certain of the relation of geography to color pattern, but the available evidence suggests a northwest to southeast cline. Table 1. Color of Apical Sternite of Female Locality Number of S pccimens Largely yellow apically Tip yellow Tip yellow all across on sides California1 2 Wellton, Ariz. 1 Rodeo, N. Mexico 2 7 Lordsburg, N. Mexico 4 Albuquerque, N. Mexico 1 Mexcala, Guerrero 2 Pt. Isabel, Texas 1 42 Table II. Color of Mesosternum of Male Locality Number of specimens Almost Black spots Posterior .6-. 8 Sternum and pleura wholly posteriorly black, some black except yellow on pleura anterior border Tucson, Ariz." 2 Rodeo, N. Mex. 9 Lordsburg, N. Mex. 1 Mesilla, N. Mex.3 Pt. Isabel, Tex. 1 9 24 5 Cotulla, Tex4 2 1 1 Paratypes of u-scripta, bearing no further locality data. These specimens otherwise have the yellow maculations generally reduced as compared to Arizona specimens, and also have a large black spot on the clypeus. Types of u-scripta Par a type of arcuata 4 Type and paratypes of arcuata Observations on Colony at Port Isabel, Texas This colony was first discovered June 23-25, 1956, but at this time it was evident that the wasps had nearly completed their activities for 1960] Evans — Bembix u-scripta 49 the season. Only a few females with badly worn wings and mandibles were still active, and even these ceased activity after June 25 (Evans 1957). I returned to the area May 8-1 1, 1958, and found the wasps at the peak of their activity. C. S. Lin revisited the area June 3-7 of the same year and found that the males had disappeared and that nest- ing of the females was more advanced. Thus in this area the wasps first become active in early May (probably in late April) and the females remain active for six to eight weeks. The species is certainly univoltine, as cocoons collected June 23, 1956, gave rise to adult males and females in late March, 1957 (they were overwintered at room temperature at Ithaca, N. Y.). The colony was located on the protected beach of Laguna Madre about five miles west of Port Isabel. The soil here is a rather heavy, dark sand containing pieces of shells and bits of organic matter (chiefly bits of decayed seaweed and beach halophytes). This beach is occasion- ally swept by waves from Laguna Madre (which is salt), especially during the fall and winter, but there is little tide action. On certain parts of the beach there are extensive patches of low vegetation, chiefly Salicornia , certain composites, and various grasses. The wasps oc- curred only in bare places on the middle of the beach, that is, from about 8 to 15 meters above the high water mark hut below the banks at the upper edge of the beach. Behavior of males. — Males were observed only during early May, 1958, but at that time they were very abundant. Each morning we entered the area about 0800, and at this time the males were already engaged in their “sun dance”. Each male flew rapidly in loops, figure eights, and irregular patterns only 2-6 cm. above the sand surface, much as in Bembix cinerea (Evans 1957). By 1030 some reduction of males could be noted, and by 1115 each day the last male had dis- appeared. By 1500 a few males were once again active, and from 1600-1900 each day there was a second though somewhat less popu- lous sun dance. The greater part of this sun dance occurred in a small area somewhat apart from the major nesting area of the females, although males often flew briefly over various parts of the beach before rejoining the sun dance. During the middle of the day, and also at night, the males enter short, oblique burrows in the sand and throw up a small barrier of sand behind them. These burrows tend to be grouped in small clus- ters. They are dug by the males, but the males usually re-enter an old burrow rather than digging a new one each day. 50 Psyche [September During the sun dance the males fly, at least some of the time, with their middle and hind legs hanging down. When they approach a fe- male they attempt to descend upon her from above and grasp her with the legs. During this time the pair rise a meter or two in the air, then, if the female is receptive, descend to the ground, where copulation occurs. During mating, the male rests on top of the female, often buzzing his wings (perhaps to maintain balance), while the female re- mains completely motionless, her wings being clamped in place by the middle legs of the male. The spines of the middle femora of the male apparently fit over the wing-bases of the female, while the arcuate middle tarsi of the male appear to fit over the coxae of the female. The front legs of the male extend rigidly forward and the hind legs rigidly backward during copulation ; neither pair is in con- tact with the female or the substrate. The male genitalia are of course extruded and work up and down rhythmically until they gain entry into the genital chamber of the female. The longest copulation observed lasted two minutes. Nesting behavior of females. — Most of the females were concen- trated in one large bare area on the beach. In this area there were at least 100 nests, many of them only a few centimeters apart. Nests on the periphery of the colony tended to be more widely separated. Here and there along the beach there were other smaller groups of nests, so that the total may have approximated 200. Most new nests appear to be started in the morning (0900-1030). The female is able to dig about 10 cm. an hour, but because of fre- quent periods of inactivity, especially at midday, the greater part of a day is generally required to complete a nest. Much use is made of the mandibles in loosening the soil, and this results in much wear of the mandibles as the season progresses. The soil which is loosened is scraped back with the front legs in the usual manner of wasps of this genus. When the burrow is a few centimeters deep, the wasp allows the soil to clog the entrance, sometimes for long periods, before finally coming out and scraping it back from the entrance. The digging movements appear rather slow as compared to other species of the genus. As digging progresses, a large mound of sand accumulates behind the entrance. This mound measures, on the average, about 18 cm. long, 10 cm. wide, and 1.5 cm. high. When the nest is completed, some individuals level this mound. The leveling movements aie very characteristic and unlike those of any other known Bembix. The fe- 1960] Evans — Bembix u-scripta 5i male (without closing the nest entrance) backs out to the middle of the mound of sand, then walks forward in a straight line scraping sand backward. When she arrives at the nest entrance, she turns to one side at a 45-90° angle and continues for another 1-3 centimeters, still scraping sand, which now falls over or slightly beyond the nest entrance. She then flies back to the center of the mound and repeats Fig. 1 — Diagram of leveling movements of Bembix u-scripta. In a is shown the completed mound of earth with the open (blackened) nest en- trance at one end. In b three typical routes of a female leveling the mound are shown; dashed lines indicate movement by flight. In c is shown the final product, with the nest entrance covered (indicated by dashed line) and mound well removed from entrance. the performance. After a time this results in the soil immediately in front of and around the nest entrance being well scattered and flat- tened. However, the extremity of the mound of soil has not been touched and has in fact been added to. Thus the net result of the leveling movements is to move the bulk of the accumulated soil from the nest entrance to a position well removed from it (by about 8-12 cm.) (fig. 1). In this colony relatively few individuals leveled the mound. We estimated that about 25% leveled, while some others exhibited weak and indefinite leveling movements. The remaining members of the colony left the mound of earth completely intact. As additional cells are added to the nests, some additional soil is added to the mound; 52 Psyche [September as a result nests containing several cells often have a very large mound of earth (as much as 3 cm. high). It was our impression that indivi- duals that level following completion of the initial cell also level after each additional cell, but our data on this point are weak. When the nest or new cell is completed, and leveling is completed if this is to occur, the female flies off to obtain her first fly, on which the egg is laid. This invariably occurs in the late afternoon or early evening, and after oviposition the female closes the nest from the in- side for the night. Thereafter the nest entrance is left closed from the inside when the female is inside for long periods of time (at night and during much of the day). During these periods the cell is also closed off from the burrow by a small barrier of sand. The female remains in the bottom of the burrow just outside this barrier or “inner closure”. During periods of provisioning, however, both inner and outer close- ures are removed. Nest entrances are occasionally found to be open for brief periods in the morning, probably while the female is out taking nectar from flowers. In this area no wasps were actually ob- served visiting flowers. Final closure of the nest is not especially distinctive and has already been described (Evans 1957) * Description of nest. — - The burrow measures about 15 mm. in diameter, generally slightly more than this near the entrance. It enters the soil at a 45-70° angle with the horizontal but tends to level off as it approaches the horizontal cell. The burrow is often straight but may have a weak to strong lateral curve part way down. Measure- ments of 36 nests showed the burrow length to vary from 19 to 40 cm., with a mean of 30 cm. Depth of the bottom of the cell, measured vertically from the soil surface, varied from 10 to 22 cm., with a mean of 15 cm. Cells measured about 18 mm. in diameter and from 30 to 40 mm. in length. Of 16 nests dug out May 8-1 1, 1958, 14 contained one cell and two contained two cells; none of these nests had yet received the final closure and hence all might have eventually had additional cells. Of 20 nests dug out by C. S. Lin June 4-7 of the same year, 1 1 contained one cell, 8 contained two, and one contained three. Only a few of these (including the three-celled nest) had received the final closure. The several nests dug out in late June, 1956, were nearly all multi- cellular (Evans 1957). Two nests were tricellular; one of these was still active while the other was receiving the final closure when dug out. Two other nests appeared to have four cells and another five. However, in these cases all the cells contained cocoons and the burrows 1960] Evans — Bembix u- script a 53 leading to them had been filled and hence could not be traced accurate- ly. Because of the close proximity of nests in this colony, it is im- possible to be certain that all of these cells belonged to the nest in question. The evidence now seems strong that the nest is usually closed after the completion of only two or three cells. It is not im- probable that one-celled nests also occur, perhaps also four or five- celled nests rarely. Two or three-celled nests are characteristic of Bembix nubilipennis (unpublished observations), while B. cinerea and B. belfragei characteristically make one or two-celled nests (Evans 1957). Fig. 2. — Typical three-celled nest of Bembix u-scripta (our field note no. 1635, Rodeo, N. Mex.). Cell a contained a completed and hardened cocoon, so was obviously the first cell of the nest. Cell b contained a fresh, still soft cocoon; cell c was a new cell which was empty when the nest was dug out. The figure on the left shows the nest in lateral view, compressed to a single plane, while the sketch on the right shows the proper relationship of the cells in a horizontal plane. I he cells of a given nest are normally separated by several centi- meters of soil. After the first cell is fully provisioned, the bottom .3 to .5 of the burrow is packed tightly with sand. 1'he wasp then con- structs another section of burrow of about equal length at roughly a 90° angle with the main burrow and builds the second cell at the end of it. The third cell-burrow and cell are normally constructed from the opposite side of the main burrow (fig. 2). Provisioning. — During the first two days of study, we were unable to find females provisioning their nests even though the nests which we dug out usually contained fresh flies. We left the area on these days at 1600-1700, a time when most digger wasps have completed 54 Psyche [September their activities for the day. On the third day we noted a few females provisioning their nests about 1600, and remained in the area until 1 9 1 5 • We discovered that there was a great emergence of females from their nests just before sunset. During the two-day period of observation, the first female carrying prey was seen at 1520, the last at 1905, with the great bulk of the provisioning occurring between 1700 and 1900. Sunset was at about 1830. Some females and males were still active at 1 9 1 5 > when it was too dark for us to see them well, but apparently these individuals were merely digging in for the night. In order to determine what types of dies the wasps were taking at this unusual hour, we took 97 dies from females or from the nests. These dies represented 33 species of nine families; that is, there was an average of only three dies of any one kind. The most numerous dies were those of the genus Acrosticta (Otitidae) (26 examples). Bombyliidae, Asilidae, and Tachinidae were represented in roughly equal numbers, each family by several species. A complete list is pre- sented in Table III. Although many species of Berubix are relatively unselective of their dies, I have never encountered a more remarkable diversity of prey in any one locality. Most of the dies used were rel- atively small, and a great many of them were used per cell ( about 40) . However, some relatively large dies were found in some nests. It required very little study to determine that virtually all Diptera had ceased activity by the time the Bembix began to provision in num- bers. Some of the dies (e.g., the otitids, and some of the bombyliids and sarcophagids) were seen commonly on the beach during the middle of the day, but in the evening they had disappeared. 1 he Bembix ap- parently hunt widely in the vegetation and take virtually any “sleep- ing” dies within a certain size range which they encounter there. Females carrying dies enter the nesting area with a rather loud buz- zing only 10-20 centimeters high. They plunge quickly into the nest entrance which, as noted above, is never closed during provisioning. In less than a minute, they re-emerge from the burrow and quickly take wing. Several instances were observed of females attempting to steal dies from one another. Oviposition and development. — The first dy is placed deep in the cell on its back, head-in, with one wing somewhat extended. The egg is laid erect, glued to the side of the dy near the base of the extended wing, or actually glued to the wing-base. One instance of abnormal oviposition was observed. In this case the dy (Acrosticta sp.) was actually smaller than the egg and was dorsum-up, with the egg of the wasp glued to the base of the abdomen and extending obliquely for- 1960] Evans — Bernbix u-scripta 55 ward. The egg of u-scripta measures about 4.5 mm. long. The egg hatches in about two days and the small larva remains at- tached to the fly pedestal by a glutinous thread for several days. Pro- visioning is fully progressive; that is, flies are provided each day up until the larva is full-grown and nearly ready to spin its cocoon. The wasp does not clear the fly remains from the cell, and they mere- ly accumulate in a mat on the bottom of the cell. Apparently about a week is required for completion of larval development. TABLE III. PREY OF BEMBIX U-SCRIPTA AT PORT ISABEL, TEXAS Family and species of fly Number taken Stratiomyidae Hermetia aurata Bell 1 Tabanidae Tab anus texanus Hine 1 Bombyliidae Aphoebantus sp. nr. hirsutus Coq. 6 Lordotus gibbus gibbus Lw. 5 L. gibbus striatus Painter 3 Phthiria sp. 1 Poecilanthrax lucifera Fabr. 3 Villa parvicornis Lw. 2 Asilidae Erax cressoni Hine 2 E. tuberculatus Coq. 4 Pleisosomma unicolor Lw. 1 Psilocurus modestus Will. 2 P. puellus Bromley 2 Stenopogon ebyi Bromley 6 Syrphidae Volucella fraudulcnta Will. 1 V. unipunctata Curran 3 V. sp. 1 Otitidae A crosticta mexicana Cole 15 A. sp. 1 1 Muscidae Limnop/iora sp. 1 Mydaea sp. 1 Phyllogastcr cordyluroides Stein 5 Sarcophagidae Sarco phaga johnsoni Aid. 4 S. sp. 1 Senotainia kansensis Tns. 1 Tachinidae Archytas marmoratus Tns. 1 CAiaetogaedia sp. nr. analis Wulp J Goniochaeta plagioides Tns. 1 Opelousia obscura Tns. 1 Phorocera tachinomoides Tns. 1 Promasiphya confusa Aid. 1 T dchinophyto sp. nr. vanderwulpi Tns. 3 Xenoppia monela Rnh. 7 56 Psyche [September Observations on Colony at Rodeo, New Mexico 1 his small colony of Bembix u-scripta was discovered on August 28, 1959, in open desert at about 4000 feet elevation one mile north of the town of Rodeo, in extreme southwestern New Mexico. The area in question has only a few shrubs and small trees (chiefly mesquites and yuccas), but following the summer rains it is densely covered with composites and other flowering annuals, Russian thistle, and grasses. Here and there are small places devoid of vegetation. In some of these the ground is flat and the soil rather compacted, while in others the soil is loose, sandy, and somewhat blown-out or piled up in small dunes no more than a meter or two high. One of these small dunes contained about six nests of Bembix u-scripta , while two other nests were found in similar places not far distant. A colony of A philanthops haigi was located in the same dune with most of the Bembix nests, the nests of the two wasps intermingled. A few specimens of Bembix sayi and B. Similans also nested within the total area inhabited by B. u- scripta. Many things about this situation were strikingly different from the Port Isabel location. The season was of course different, though August is properly considered “spring” in this area, as the summer rains occur in July and August and result in a great flowering of the desert and in the appearance of many Hymenoptera. The soil here was of finer texture, more friable, and apparently drier, although oc- casionally moistened temporarily by showers. No tests were made to determine whether the soil was of comparable salinity, but surely it was not, as this is not an area of notable internal drainage such as oc- cur in some parts of the Southwest. The total population of Bembix • u-scripta in this area may have been no more than 30 to 40 individuals of both sexes. Consequently, in spite of the longer period of study (August 28- September 12), my data are more fragmentary. However, I was especially alert for re- semblances to and differences from the Texas colony, and the follow- ing account stresses these. Behavior of males. — The first males were observed August 21 fly- ing over certain bare, sandy patches, although no females were dis- covered until a week later. The flight of the males was extremely rapid and the pattern of flight very irregular; it was only after many failures that I succeeded in taking one for identification. There were scarcely enough males for a typical “sun dance”, though now and then two or three would fly about the same small area and make contact 1960] Evans — Bembix n-scripta 57 with one another in flight briefly. Males were generally active 0800- 1130 and again, in lesser numbers, 1600- 1800. Several were seen visiting the dowers of Gaillardia pulchella for nectar, as were several females. At 1630 on September 8 a pair was seen in copula on these flowers. I hey remained together for two minutes, then separated briefly and once again came together. The details of copulation were essentially as described for the Texas colony. Nesting behavior of females. — The nests in this area tended to be well separated, the closest nests being about half a meter apart. New nests were started in the morning and completed in the late after- noon; during much of the day the entrance to such a nest would be closed with a plug of sand thrown up from the inside, where the fe- male was presumably resting or digging only intermittently. During the more active periods of digging the wasp now and then comes out and clears the sand from the entrance. Eventually a fairly large mound of sand accumulates outside the entrance. When the nest is completed, the female comes out (without closing the entrance behind her) and commences to level this mound of sand. The leveling move- ments were found to be precisely the same as in the Texas colony (fig. 1). They result in the mound being not so much leveled as spread out and moved slightly farther from the nest entrance. Follow- ing leveling, the 10 cm. directly in front of the entrance are typically smooth, with beyond this the mound, measuring about 20 cm. long by 18 cm. wide. Leveling was observed several times and little variation was noted; always it occurred in late afternoon and required 15-25 minutes. Following completion of a new cell in an old nest, little addi- tional sand is usually thrown out, but such sand as accumulates is treated in the same manner. Also, when a female emerges from her nest in the late afternoon and is about to begin bringing in prey, she clears the entrance of sand and proceeds to level this sand briefly in the same manner. All individuals observed exhibited these leveling movements. The general structure of the nests in this area was very much as described for the Texas colony. However, they were slightly deeper. In the eight nests which were dug out, burrow length varied from 30 to 42 cm. (mean 35 cm.), cell depth from 16 to 23 cm. (mean 18 cm.). Five of the eight nests had but one cell, while two had twro and one had three (fig. 2). None of these nests had received the final closure when I dug them out, so all might eventually have had more cells. However, since I saw several females digging new nests toward 58 Psyche [September the end of the period of observation, I doubt if the number of cells per nest is normally more than two or three. Nests dug out when the female was not provisioning were invariably found to have two closures, one at the entrance and one just outside the cell, with the female remaining just outside the inner closure. During provisioning, and also when the female is out taking nectar from flowers, the outer closure is not maintained, though the inner closure is normally maintained if the cell contains an egg or small larva. Provisioning. — All provisioning occurred in the late afternoon and early evening, as in the Texas colony. During the period of observa- tion, sunset occurred at about 1820-1830. However, the area in ques- tion was located directly east of the Chiricahua Mountains; the sun disappeared behind these shortly after 1800, leaving the area in deep shadow. The first female carrying prey was observed at 1600, the last at 1842, with the bulk of the provisioning occurring between 1730 and 1830. September 10 was a cloudy day, and several females were out of thenests earlier than usual (by 1430) ; however, none were seen with prey until about 1 700. Several females were observed hunting flies. They would fly about tall herbs (especially Russian thistle, Salsola kali tenuifolia ) and actually fly in amongst the branches. Evidently they were searching for flies resting on the vegetation, for at this hour no flies were active. I was able to obtain only a few records of prey (Table IV). The wasps normally provide only a few fresh flies each evening, and nests dug out in the morning usually have few intact flies if any. It ap- peared that Bombyliidae provided the major prey in this area. Fe- males provision very rapidly, entering the nests quickly and remaining inside only 15-30 seconds each time. One female was observed to bring in eleven flies in one hour (1720-1820). TABLE IV. PREY OF BEMBIX U-SCR1PTA AT RODEO, NEW MEXICO Family and species of fly Number taken Therevidae C bromolepida pruinosa (Coq. ) 1 Bombyliidae Phthiria sulphur ea Loew 2 Villa (Chrys anthrax) sp. 1 V. (Villa) salebrosus Painter 1 Sarcophagidae Sarcophaga sp. 1 Tachinidae Olenochaeta kansensis Tns. 1 Phorocera sp. 1 1960] Evans — Bembix u-scripta 59 Oviposition and development are as described for the Texas colony. Invariably provisioning occurs until the larva is nearly ready to spin its cocoon. Summary and Discussion Despite the difference in season and ecological situation, the mem- bers of the two colonies studied exhibited striking resemblances in nesting behavior. The slightly deeper nests of the New Mexico colony were doubtless correlated with the lighter texture of the soil. As a general rule, with reference to populations of a single species or closely related species, wasps nesting in looser soil tend to make deeper nests. In this instance the difference in nest depth is slight, and it would be rash to claim that it is genetically determined. The fact that the New Mexico colony was much smaller and had the nests much more widely scattered may merely mean that this area was ecologically marginal for the species or that this population had undergone a decline because of unknown physical or biological factors. The one difference which is more difficult to reason away pertains to the numbers of individuals exhibiting leveling movements. In the New Mexico colony, although the number of individuals observed was not large, all appeared to level the mound at the nest entrance in the manner characteristic of the species. In the Texas colony the majority did not, but roughly 25% of the individuals leveled in exactly the same way as the members of the New Mexico colony. It would be most interesting to study colonies from other parts of the range in this regard. It is conceivable that this percentage may be found to reflect a difference in gene frequency which varies clinally somewhat as color pattern appears to do in this species. On the other hand, one cannot be certain that even this difference is genetic; it is possible that the movements are innate but subject to threshold factors which were dependent upon the differing ecology of the two areas. On the whole one cannot help but be much impressed by the similar- ity in the nesting behavior of these two widely separated colonies as well as the several unique features in the behavior of the species. In the following paragraphs some of the more significant features of the behavior are discussed one by one. ( 1 ) The species is crepuscular, at least with respect to hunting and provisioning. Some females begin hunting 2.5-3 hours before sun- set, but the majority do not become active until about an hour before sunset. From then on until about half an hour after sunset most fe- males leave the nest entrance open and are actively provisioning. The 6o Psyche [September last females close the nest at about the time it becomes difficult for a human to observe them without artificial illumination. So far as I know this is unique among digger wasps. (2) The females hunt flies which are at rest in herbaceous vege- tation. The wasps fly in amongst the plants and presumably snatch the flies from the leaves and branches. This is in marked contrast to other Bembix, which pursue flies visiting flowers or flying about ani- mals, dung, etc. The Hies taken by u-scripta are exceedingly diverse and generally rather small for the size of the wasp. ( 3 ) The mound of earth at the nest entrance is leveled in a man- ner unlike that of any other North American Bembix , though bearing some resemblance to that of spinolae and several other species. (4) The nest is multicellular (apparently usually bicellular or tricellular) , much like that of nubilipennis. (5) The nest entrance is left open during provisioning but is at all other times closed from the inside (somewhat as in sayi). (6) The egg is laid on the first fly (as in all but a few of the more specialized species of Bembix.) (7) The cell is of simple structure and is not cleaned by the female. Of these seven characteristics, the first three are unique, the fourth and fifth unusual although not unique, the last two common to all the more generalized species of Bembix (several more characters of this nature could easily be tabulated). Thus the total picture is not unlike that obtained from a study of structure: several unusual features built into a basically primitive Bembix. The most interesting of these features pertain to the unusual time of flight of provisioning females and the fact that they take Diptera which are at rest in vegetation. The well developed ocelli probably represent part of the morphological component of this adaptive complex. Since the ancestral Bembix doubtless had ocelli of nearly normal sphecid type, it seems probable that u-scripta split off from this stock at a very early date, before the loss of ocelli became complete. This would also account for the fact that certain features of wing venation and male secondary sexual characters are more Stic tie Ha-\ike than Bembix- like. 1 he crepuscular provisioning behavior of u-scripta clearly enables the species to exploit a slightly different source of food from that of other species of the genus: a miscellany of mostly rather small flies which are snatched from their resting positions in vegetation. So far as can be judged at present, 'u-scripta is not strongly restricted ecologically ; perhaps the 1960] Evans — Bc?nbix u-scripta 6l species is able to live side-by-side with other Be?nbix by virtue of its unusual provisioning behavior. Since u-scripta is one of the rarest of North American Bembix, I would hesitate to say that its unusual adap- tations have proved outstandingly successful. A final word on the second problem, that of the specific status of arcuata and u-scripta. I find it hard to believe that the wasps which I studied at Port Isabel, Texas, and Rodeo, New Mexico, represent different species. As indicated earlier, the Texas wasps were colored much like the type of arcuata, the New Mexico wasps much like the type of u-scripta. However, they were not colored precisely like the type of u-scripta and the parameres of male genitalia were more slend- er and attenuate than they appear to be in the type of that species. The final solution must come from studies farther west, from in and around Tucson, Arizona, the type locality of u-scripta. In the mean- time I see no reason to abandon my former belief that the two names should tentatively be regarded as synonyms. Literature Cited Evans, H. E. 1957 Studies on the comparative ethology of digger wasps of the genus Bembix. Cornell Univ. Press. 248 pp. 1959 Studies on the larvae of digger wasps (Hymenoptera, Sphecidae). Part V: Conclusion. Trans. Amer. Ent. Soc., 85: 137-191. Lohrmann, E. 1948 Die Grabwespengruppe der Bembicinen. Cberschau und Stammesgeschichte. Mitt. Munchen. Ent. Ges., 34: 420-432. Parker, J. B. 1917 A revision of the bembicine wasps of America north of Mexico. Proc. U. S. Nat. Mus., 52: 1-155. 1929 A generic revision of the fossorial wasps of the tribes Stizini and Bembicini. Proc. U. S. Nat. Mus., 75 (art. 5) : 1-203. DEFENSE MECHANISMS OF ARTHROPODS. II. THE CHEMICAL AND MECHANICAL WEAPONS OF AN EARWIG.1 By T. Eisner Department of Entomology, Cornell University, Ithaca, N. Y. The European earwig, Forficula ciuricularici Linn., possesses two pairs of small sac-like glands (Vosseler, 1890) situated dorsally in the abdomen, and opening on the posterior margins of the third and fourth abdominal tergites (text fig. 1). The principal active com- ponents of their secretion have recently been identified as 2-methyl, and 2-ethyl-/)-benzoquinone ( Schildknecht and Weis, i960). The glands have always been suspected to serve in defense against pre- dators, and this suggestion, although advanced originally in lack of evidence, appears recently to have received at least indirect support, since it is now known that the quinone-containing secretion of certain other insects are, in fact, extraordinarily effective repellents (Eisner !958a, 1958b). In addition to the glands, Forficula possesses the typical dermapteran pincers, and these too have been claimed to play a defensive role (in addition to other roles, including prey capture), although the evi- dence is here again circumstantial at best (Burr, 1910; Gadeau de Kerville, 1905; Rau, 1933). The purpose of this note is to present the results of some experi- ments designed to test whether the glands and pincers are really the effective defensive devices they were presumed to be.'J The Pincers In order to facilitate close-range observation of the earwigs, they were attached to rods (by a technique used previously and already described in detail: Eisner 1958a, 1958b; Eisner et ah, 1959), and adjusted to a normal stance on the substrate as shown in Plate 7, figure 1. In this way they were prevented from scurrying about, while at the same time retaining full mobility of head, abdomen, and legs. By subjecting such animals to mild traumatic stimuli, applied locally to the body or to individual appendages by pinching with fine forceps or touching with a warm needle, it was possible to witness very clear- ly the way in which the pincers are put to a defensive use. This study was supported by Grant E-2908 of the U. S. Public Health Service. 2 Forficula auricularia is established in the U. S. A. The specimens used were taken in Lexington, Mass. 62 1960] Eisner — Deferise Mechanisms 6 3 No sooner had a stimulus been applied than the animals responded by revolving their abdomen, bringing the pincers to bear upon the region traumatized. They usually succeeded in grasping the instru- ment used for stimulation ( Plate 7, figs. 2, 3) and, by pulling and tugging, attempted to free themselves from it. They sometimes re- leased their grip momentarily, but as a rule persisted tenaciously for as long as the stimulus was maintained. The abdomen is remarkably maneuverable, being capable of bending and twisting to such an Text Figure 1. Forficula auricularia Linn, showing the four glandular- openings and the cuticular sculpturing around them on the third and fourth abdominal tergites. extent that virtually no body part other than the abdomen itself is inaccessible to the pincers. Since the abdomen responds rapidly and with precision, the pincers can be shifted accurately and without delay from one region to another. This became quite clear when the ani- mals were subjected to a rapid sequence of stimuli applied to different parts of their body. The pressure that the animals can exert with the pincers is con- siderable. From personal experience, I can attest to the claim (cited by Burr, 1910) that the sharp prongs can pierce human skin (the prongs of the male are larger and somewhat more effective than those of the female) . The Glands Adult earwigs, affixed to rods as before, were again subjected to localized stimulation, but this time they were placed on sheets of filter paper impregnated with an acidulated Kl-starch solution. This paper discolors to an intense blue-black in the presence of quinones, and can therefore be used as an appropriate indicator for the detection of glandular discharges. In Plate 8 are shown the types of pattern produced by the secre- tion. Evidently, the gland contents do not merely ooze out, — as 64 Psyche [September they do in certain millipedes and tenebrionid beetles that also dis- charge quinones (Eisner, 1961) — but rather are ejected forcibly as a spray. It is also apparent from the figures that the spray is not ejected in a fixed direction, but is actually aimed with considerable precision toward the particular region of the body subjected to stimu- lation. This finding, although new for earwigs, is by no means novel as it applies to arthropods in general : bombardier beetles, certain cockroaches, whipscorpions, and a variety of others, are all known to aim their spray (Eisner, 1958a, 1958b, 1961; Eisner et al., 1959; Eisner et al., 1961 ) . The mechanism by which the earwigs aim is intimately linked with the defensive use of their pincers, since by revolving the abdomen at its base while bringing the pincers toward the stimulus, the gland openings are automatically pointed in the proper direction. It is interesting in this context that the forceps seem to be used in pre- cedence to the glands. When a stimulus is applied, a secretory dis- charge usually does not follow at once. As a rule, the pincers are brought into action first, and then, only after their persistent use for several seconds has proven futile, is the spray finally ejected. When the stimulus applied is a more violent one, however, such as when the head or abdomen is pinched with hot broad-tipped forceps, or the animal simply seized between the fingers, the discharge may follow with little or no delay. In other arthropods with similar defensive glands, but lacking such auxiliary mechanical devices as the earwig’s pincers, even a slight traumatic disturbance usually elicits a prompt discharge (Eisner 1958a, 1958b; Eisner et al., 1959; Eisner et al., 1961). Up to six consecutive discharges could be evoked from a single earwig, but usually they produced no more than four. The amount of secretion ejected decreases progressively, until finally there are produced on the paper no more than a few spots at close range. The earliest that an earwig with depleted glands was ever tested to check on the restoration of its secretory supply was after a period of five days: by this time it was again able to discharge three times. Explanation of Plate 7 Figure 1. Earwig fastened to hook, in normal stance. Figure 2. Earwig having its right metathoracic leg pinched with forceps, responding b)' bringing its pincers to bear upon the forceps. Figure 3. Same as preceding, but the stimulus is applied to the right antenna. Psyche, 1960 Vol. 67, Plate 7 Eisner — Defense Mechanisms 66 Psyche [September Encounters with Predators ants [P ogonomyrmex badius (Latreille) ] The experiments were carried out under the same conditions as prevailed in previous comparable studies (Eisner 1958a, 1958b), the earwigs being affixed to rods and placed individually close to the nest entrance of a laboratory colony of the ant. Sheets of Kl-starch paper were placed under the earwigs. The ants at first attacked singly or in groups of two or three. No sooner had one of them grasped the earwig with its mandibles (often mere contact seemed to suffice), than the earwig responded by attempt- ing to catch the assailant with its pincers. Typically, the ant was seized and promptly removed from its hold, being released only after the abdomen had uncoiled and straightened out. The uncoiling of the abdomen is a rapid spring-like action, with the result that the ant is sometimes Hipped several centimeters away. No discharges were produced during these first attacks, all ants being successfully fought off by the use of the pincers alone. Gradu- ally, however, the activity of the ants around the nest entrance be- came considerably intensified (perhaps in response to the release of the ant’s own alarm substance; see Wilson, 1958), and within minutes the earwig was surrounded by a swarm of workers, some attempting to bite and sting it, others merely scurrying over its body. It was then that a discharge was finally produced. The result was an in- stantaneous dispersal of the entire swarm. For a few seconds there- after none of the ants ventured within the immediate vicinity of the earwig. Many showed typical grooming activities, others underwent the conspicuous dragging behavior already noticed in this ant and the related P. occidentals (Cresson) under similar circumstances involving insects which spray (Eisner 1958a, 1958b; Eisner et ah, 1961). The swarm soon closed in again, but for yet another several seconds there occurred no real attacks, the ants turning and fleeing the moment they made contact with the earwig. That this avoidance behavior may have been attributable to residual secretion remaining on the earwig’s body was suggested by the fact that a small strip of indicator paper held within millimeters from the earwig rapidly be- came discolored. Explanation of Plate 8 Figures 1, 2, 3. Three different discharges elicited by pinching respectively the end of the abdomen (figure 1), the right mesothoracic leg (figure 2), and an antenna (figure 3). (The pictures were taken shortly after discharge, and the abdomens were therefore no longer in the exact positions they had assumed at the time of spraying). Psyche, 1960 Vol. 67, Plate 8 Eisner — Defense Mechanisms 68 Psyche [September Eventually, within less than a minute, the earwig was again under assault. The attacks were at first countered by the pincers alone, but soon the swarm was dispersed as before by another discharge. Several earwigs were tested in this way, the results being similar in all cases. One earwig was left with the ants until well after all its secretion had become depleted. It was soon overrun by a swarm, and was by now entirely dependent on its pincers for defense. Under these con- ditions it became particularly clear how effective the pincers really are. During a period of 20 seconds that was timed, the earwig seized and successfully removed from its body a total of 19 attacking work- ers. This was, of course, a mere fraction of the total swarm, and the earwig was eventually bitten and stung repeatedly, and finally killed. During none of these encounters did any of the ants receive lasting noticeable injury. Neither did the earwigs — not, at least, for as long as their secretion lasted. praying mantids [Hierodula patellifera (Serville)] The fifteen earwigs that were given to the three adult female man- tids that were tested, were all caught and eaten in rapid succession, the mantids showing no signs of being affected by either the secretion or the pincers. The same species of mantid also accepts the quinone-secreting cock- roach Diploptera punctata Eschscholtz (Roth and Stay, 1958; Eisner !958b), hut it rejects the bombardier beetle Brachynus ballistarms Leconte (Eisner 1958a), the secretion of which is likely to be quin- onoid like that of its congeners ( Schildknecht, 1957). The reason for these inconsistencies in acceptability may be due to the fact that the various secretions do not contain precisely identical quinones, but also may have something to do with the special properties of the Brachynus spray, which differs from that of the others in that it is hot, and is discharged with a clearly audible detonation. [At the time I wrote on Brachynus (Eisner, 1958a), I was unaware of the thermal properties of its spray]. VERTEBRATES The animals tested were two small toads ( Hyla versicolor Le Conte), one bird [ Cyanocitta cristata (Linn.)], and one mouse ( Mus musculus Linn.). Both bird and mouse ate readily every one of the several earwigs that was offered to them (within a period of a few hours), betraying no abnormalities during the meal, or ill effects thereafter. It would be well to bear in mind, however, that before ruling out the repug- i960] Eisner — Defense Mechanisms 69 natorial effectiveness of an arthropod’s secretion against a vertebrate, long-term feeding experiments should be made, to test for any dis- criminatory tendencies that might eventually develop after a greater number of encounters, and particularly in situations where the pre- dator is also given palatable insects as an alternative choice. With Hyla the results were different. Five of the eleven earwigs offered (over a period of a few hours) were caught and swallowed, but the rest were promptly spat out (there was no apparent order in the sequence in which the earwigs were either taken or rejected). Whether rejection was on the basis of the secretion alone, or was also attributable to trauma induced by the pincers, could not be deter- mined. That the secretion may in itself be repellent is suggested by the fact that benzoquinones are strongly irritating when applied topic- ally to an amphibian : a mere sprinkling of a few crystals of p- benzo- quinone on Hyla invariably elicits a prompt scratch reflex. The pin- cers may also be of importance, however. In one instance a frog had difficulty rejecting an earwig, which was seen to have become firmly clamped to its tongue, and which was not removed until the frog used its forelegs to brush it away. Acknowledgements : I am indebted to Professor Carrol M. Williams of Harvard University, and also to his family, for having collected the earwigs for me, and to Professor Edward O. Wilson for use of his Pogonomyrmex colonies at Harvard. Thanks are also due Profes- sor Kenneth D. Roeder, Tufts University, who made the mantids available at his laboratory, and Dr. J. A. G. Rehn who identified them. Dr. Hermann Schildknecht, University of Erlangen, Germany, was kind enough to let me see his manuscript (Schildknecht and Weis, i960) before its appearance in print. References Cited Burr, M., 1910, Dermaptera. In: The Fauna of British India. Taylor and Francis, London. Eisner, T., 1958a, The protective role of the spray mechanism of the bom- bardier beetle, Brachynus ballistarius Lee. J. ins. Physiol. 2: 215-220. 1958b, Spray mechanism of the cockroach Diploptera punctata. Science 128: 148-149. 1961, The effectiveness of arthropod defensive secretions. Proc. Xlth Int. Congr. Entom. (in press). Eisner, T., F. A. McKittrick, and R. Payne, 1959, Defense sprays of roaches. Pest Control 27: 11-12, 44-45. Eisner, T., J. Meinwald, A. Monro, and R. Ghent, 1961, Defense mechan- isms of arthropods. I. The composition and function of the spray of the whipscorpion, Mastigoproctus giganteus (Lucas) (Arachnida: Pedipal- pida). J. ins. Physiol, (in press) 70 Psyche [September Gadeau de Kerville, H., 1905, Note sur les functions de la pince des insectes Orthopteres de la famiile des Forficulides. Bull. Soc. Zool. France 30: 53- 63. Rau, P., 1933, The Jungle Bees and Wasps of Barro Colorado Island. Von Hoffman Press, St. Louis, Mo. Roth, L. M., and B. Stay, 1958, The occurrence of />tfra-quinones in some arthropods, with emphasis on the quinone-secreting tracheal glands of Diploptera punctata (Blattaria). J. ins. Physiol. 1: 305-318. Schii.dk necht, H., 1957, Zur Chemie des Bombardierkafers. Angew. Chem. 69: 62-63. Schildknecht, H., and K. H. Weis, 1960, VI. Mitteilung liber Insektenab- wehrstoffe. Zur Kenntnis des Pygidialdriisensekretes vom gemeinen Ohr- wurm, Forficula auricularia. Zeitschr. f. Naturforsch. 15: 755-757. Vosseler, J., 1890, Die Stinkdrusen der Forficuliden. Arch. mikr. Anat. 36: 565-578. Wilson, E. O., 1958, A chemical releaser of alarm and digging behavior in the ant Pogonomyrmcx had'ius ( La treille ) . Psyche 65: 41-51. A TRIASSIC ODONATE FROM ARGENTINA By F. M. Carpenter Harvard University The fossil insect described in this paper was collected in 1958 in the precordilleran region of Mendoza in Argentina by the joint ex- pedition of the Museum of Comparative Zoology and the Museo Argentino de Ciencias Naturales of Buenos Aires. The specimen was turned over to me for study by a member of the expedition, Professor Bryan Patterson. The rock containing the insect is a nearly white shale, with definite bedding and numerous plant fragments. The insect consists of the distal part of a wing, very clearly preserved. About 6 cm. away from this wing there is another specimen, consisting of a small, distal fragment of a wing. The larger specimen is clearly odonate but cer- tain structural details eliminate it from all described families of the order; because of our lack of knowledge of the proximal part of the wing, including the arculus region, I am placing this species in In- certae Sedis , in preference to establishing a new family on so few de- tails. The smaller wing fragment can be interpreted best by compari- son with the larger specimen, as will be noted below. Order Odonata Family: Incertae Sedis Genus Triassothemisj new genus Pterostigma well formed, elongate; nodus incomplete, without a costal indentation at the junction of the subcosta; nodus remote from the base of the wing: about 6 postnodals between nodus and ptero- stigma. The genus appears to be related to the suborder Archizygop- tera, which has been found in the Triassic of Australia and Jurassic of Europe and Asia (Turkestan). Type species: Triassothemis mendozensis, new species Triassothemis mendozensis , n. sp. Text figure 1 A, plate 9. Wing: length (as preserved), 25 mm.; width, 8.5 mm. Ptero- stigma about four times as long as wide, distinctly pigmented; costal border along the pterostigma somewhat thickened. Nodus incipient; subcosta distinctly thickened as it meets the costa; a short cross-vein between the end of the subcosta and Ri, this cross-vein being almost aligned with the cross-vein below it (extending from Ri to R2). R3 separating from R2 at a point not much distal of the nodus; R2 and 71 72 Psyche [September R3 widely divergent distally, with four rows of cells between them. The origin of R4 + 5 from R2 + 3 is not shown in the specimen but it is clearly proximal of the level of the nodus. MA and R4 + 5 are markedly divergent distally. IA is not recognizable in the part of the wing that is preserved. Cross-veins are numerous and are dis- tributed as shown in figure ia. Holotype: No. 18040, in Museo Argentino de Ciencias Naturales, Buenos Aires; collected in the Estratos de Potrerillos, near Agua de las Avispas, along the south slope of Cerro Cacheuta, Argentina. The formation is of Triassic age (see Romer, i960, for additional geological data). The preserved portion of the wing is very clear (see plate 9) and the venational convexities and concavities are dis- tinct. Text-figure IA. Drawing of Triassothemis mendozensis, n. sp., from the Triassic of Argentina (holotype). Text-figure IB. Drawing of wing fragment associated with above speci- men. Sc, subcosta : n, nodus ; Rl, radius ; IR2, R2, R3, IR3, R4+ 5, branches of radial sector; MA, anterior media; CuP, posterior cubitus; — , concave veins; +, convex veins; pt, pterostigma. The most striking feature of this wing is the nature and location of the nodus. In its form, the nodus is like that of the Archizygoptera Psyche, 1960 Vol. 67, Plate 9 Photograph of holotype of Triassothcmis mc?idozensis , n. sp. X5 74 Psyche [September but its position is very different. In the Archizygoptera the nodus is very remote from the pterostigma and close to the base of the wing, the subcosta being very short. In T riassothemis , the nodus is close to the pterostigma and probably at about the middle of the complete wing. The nodal area is so clearly preserved in the specimen of Tri- assothemis that there can be no doubt about its structure; although the subcosta has been broken away just before its termination, it is present on each side of this missing section. The distal location of the nodus apparently eliminates T riassothemis from the Archizygop- tera, as the suborder is presently understood. Another peculiarity of the wing of T riassothemis is the widening of the area between Ri and the costal margin of the wing, just beyond the nodus. That this is not a distortion is shown by the normal location and distribution of the cross-veins and the other veins in the anterior part of the wing. The pterostigma is like that of the Archizygoptera and the several branches of the radial sector have the divergent arrangements char- acteristic of that suborder, although the divergences are not quite so marked. The absence of a distinct anal vein (at least in the part of the wing which is preserved) is also suggestive of the Archizygoptera. In most respects, therefore, Triass ot he mis possesses many of the char- acteristics which we find in the wings of the Archizygoptera but the location of the nodus is very different from that of all of the genera of this suborder so far known. The small apical fragment of a wing, which is located on the same piece of the rock as T riassothemis, resembles the latter in so far as the known parts of the venation are concerned (Figure ib). Ri, R2, IR2 are arranged about as in mendozensis but there is a clear indication that the next vein, R3, would be somewhat more remote from IR2 than it is in the specimen of mendozensis. The pterostigma is like that of mendozensis so far as its distal part is concerned, but it appears to extend proximally the full length of the preserved part of the wing, as shown in figure ib. It seems very likely that this small wing fragment represents an odonate closely related to T riassothemis mendozensis ; in fact, since we do not know anything about the dif- ferences between the fore and hind wings of the Archizygoptera, it is possible that the fragment is part of another wing of the type of mendozensis. The discovery of this piece of Triassic rock containing two fossil insects, even though they are in fragmentary condition, points up the possible significance of Triassic strata in the Mendoza region of Argentina as a source of extensive collections of insects. Wieland 1960] Carpenter — Triassic O donate 75 1925, 1926) has described two other Triassic insects collected in a Triassic deposit very near Cerro Cacheuta, one of these being a homopteron (Tipuloidea rhaetica) and the other a possible trichopteron (N ecrotaulius (?) affinis) . Both of the specimens on which these species were based are preserved on one piece of rock. In addition, Kurtz in his atlas of fossil plants from Argentina (1921) includes three sketches of insects which were found in the Triassic beds of the Mendoza area; these are stated in the caption of his plate 7 to be hymenopterous wings, although they are most certainly not rep- resentatives of that order. Also, one of the Triassic specimens which Kurtz identified as a plant (Beira argentina) and figured on his plate 22, fig. 336, is actually part of a wing of an insect, subsequently desig- nated by Cockerell (in Wieland, 1926) as Elcana (?) argentina. It seems clear from this casual collecting of insects in the Triassic deposits of the Mendoza area that further exploration of these strata for in- sects is very desirable. Literature Cited Kurtz, F. 1921. Atlas de las Plantas Fosiles de la Republica Argentina. Actas de la Acad. Nac. de Ciencias en Cordoba 7: 133-153; plates 1-27. Romer, A. S. 1960 . Vertebrate-bearing Continental Triassic Strata in Men- doza Region, Argentina. Bull. Geol. Soc. Amer. 71 (9): 1279-1294. Wieland, G. R. 1925. Rhaetic Crane Flies from South America. Amer. Journ. Sci. 9 : 21-28. Wieland, G. R. 1926. South American Fossil Insect Discovery. Amer. Journ. Sci. 12: 130-135. ON THE TRUE NATURE OF THE AZYGETHIDAE (CHILOPODA: GEOPHILOMORPHA)* By R. E. Crabill, jr. Smithsonian Institution, U. S. National Museum, Washington D. C. The family Azygethidae was proposed for the reception of a single genus and species, Azygethus at opus, by R. V. Chamberlin in his 1920 study1 of the Australian region’s myriopod fauna. The new group’s suprageneric rank was defended really on two grounds. The ultimate pedal segment reportedly had normal pleural sclerites, but even more remarkably this segment was said to bear a pair of spiracles. If the members of the genus did indeed all normally possess ultimate pedal segment spiracles, then their allocation to a new family would surely be justified, for they would thereby differ, not only from all other Geophilomorpha, but even from all other centipedes. This character was so extraordinary that, sixteen years later, At- tems expressed reservations as to its authenticity, and, regretting the total lack of figures, he urged that the specimen be re-examined. Nevertheless, in 19262 he did include Azygethidae in his treatment clearly implying his suspicion that the critical character might have been misrepresented originally. And there the matter has remained until the present. No subse- quent specimens of at opus have ever been discussed. No new species have ever been referred to the family. The original types have never been re-examined. In short, no corroborative evidence has ever been adduced in support of Chamberlin’s original interpretation. Furthermore, if we were to discount, for the moment, the spiracles and pleurites of the ultimate pedal segment and then attempt to imagine to what other family and genus the species could belong, our endeavor would prove fruitless. There are so many critical errors in the original description to lead one astray, that even a man of At- *This study was undertaken with the aid of a grant from the National Science Foundation and is published with the aid of a grant from the Museum of Comparative Zoology at Harvard College. The Myriopoda of the Australian Region, Bull. Mus. Comp. Zool. Harvard, 64(1) : 32, (1920). 2Handbuch der Zoologie, Kukenthal and Krumbach, Bd. 4: 366, (1926). of families. Two years later he failed even to mention the name3, but 'The Myriopoda of South Africa, Ann. South African Museum, 26, (1928). in his great monograph of 19294 he referred to the family again, 4Geophi!omorpha, in Das Tierreich, Lief. 52: 347, (1929). 76 1960] Crab ill — A zygethidae 77 terns’ broad experience and at times almost clairvoyant insight could not possibly have guessed to which of the existing genera and families at op us might be referrable. Thanks to the hospitality of Dr. Herbert W. Levi, who is in charge of the Arachnida and Myriapoda at Harvard’s Museum of Comparative Zoology, I have been able to study the holotype and paratype. Both are females that were collected by W. M. Mann at Levuka on Fiji. Without the slightest doubt in my mind, both are referrable to the oryid Orphnaeus brevilabiatus (Newport), which is probably the most widespread and common geophilomorph of the world’s tropics. Let us examine certain parts of the original description in light of what the type specimens themselves disclose. I shall not discuss those features that are accurately, or essentially accurately, described therein. The mandible is said to have “a single pectinate lamella”. At lower magnifications this appears superficially to be the case; however, optimal preparation and observation reveal the mandibles unquestion- ably to be those of an oryid; they are not geophiliform. The pectinate lamellae are simply pressed tightly against the distal end of the man- dible, giving it, at first sight, a geophiliform appearance. The “coxae”, i. e. the coxosternal sides, of the first maxillae are reportedly “wholly discrete”, that is, totally separated. If the reader will examine figure i, plate 10, he will see that the coxosternum is continuous and that there is not the slightest division at point E. \ he two medial processes or lobes, D, are of course discrete, as they al- ways are. Perhaps Dr. Chamberlin confused the two in preparing his original analysis. Ventral pores are said to be absent, but ventral pores are present and, in aggregate, form the patterns that are so distinctive of brevila- biatus. “Last pediferous segment with coxae distinct from the pleurae. . . Precisely what Chamberlin meant here is unclear; however, his use of the world pleurae, which is plural, implies a reference to pleural sclerites rather than to lateral body wall or membrane. In fact, be- tween the leg base and tergite there is a weakly sclerotized plate-like protuberance which appears to be an out-folding of the lateral body wall (plate 10, fig. 2, F). This same structure is to be seen in speci- mens of O. brevilabiatus ; it is more pronounced in some than others. Probably it represents an abortive paratergite, a serial homologue of the more anterior, typical paratergites. In any event, the statement 78 Psyche [September on p. 32, that “the pleural plates of the segment [are] normal” is quite misleading. Finally, the ultimate pedal segment was reported to have a pair of spiracles, and in fact it has such spiracles (fig. 2, F). At the same time, there are mitigating circumstances. First, these spiracles occur only on the holotype : the paratype, otherwise identical with it, lacks them altogether. Secondly, the nature of the holotype’s spiracles clearly suggests them to be anomalous, in this case probably ontogene- tic freaks. Both are abortive, the left one more so than the right, and there is no evidence that either was ever functional. Both evidently lack a connection with the tracheal chain linking the more anterior, normal spiracles. Such spiracular anomalies are not unknown. Indeed, I have report- ed having found one in a cryptopid centipede, Scolopocryptops s ex- spin osa (Say)5. In this case anomalous spiracles were discovered on the 7th pedal segment, which normally lacks them, although in a related genus, Dinocryptops , the 7th segment normally has them. Thereby the old mystery surrounding the apocryphal family Azyge- thidae appears to be resolved. Azygethus atopus Chamberlin is a junior synonym of Orphnaeus hrevilahiatus (Newport), and Azyge- thidae falls as a junior synonym of Oryidae. In closing I cannot help but wonder what other chilopod names owe their raison d'etre to noth- ing more than to such sporadic, anomalous characters, and how long they will continue to obscure the real fabric of the system before their paroles are terminated. 5On the reappearance of a possible ancestral characteristic in a modern chilopod, Bull. Brooklyn Ent. Soc., 50 (5): 133-136, (1955). Explanation of Plate 10 Azygethus atopus Chamberlin figure 1. Paratype. First maxillae: left two-thirds, with adjacent second maxillae, the setae deleted. A — lappet of coxosternum. B— -lappet of telopodite. C telopodite. D right medial lobe. E = coxosternum. F = hid- den continuation of coxosternum concealed beneath anterior edge of second maxillary coxosternum. Figure 2. Holotype. Posterior-most body segments: right side tilted slightly downward, setae deleted. A — tergum covering the postpedal seg- ments. B — tergite of ultimate pedal segment. C — Pretergite of ultimate pedal segment. D=Tergite of penultimate pedal segment. E = right ultimate leg. F = Spiracle and plate-like bulge of ultimate pedal segment. G = Coxo- pleuron. H — Parapretergite. I = Stigmopleurite of penultimate pedal seg- ment. J = Paratergite of penultimate pedal segment. Psyche, 1960 Vol. 67, Plate 10 A 2 Crabill — Azygethidae CAMBRIDGE ENTOMOLOGICAL CLUB A regular meeting of the Club is held on the second Tuesday of each month October through May at 8:00 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 shows the head of the worker of Myrmotcras karnyi Gregg. The original, drawn by Dr. R. E. Gregg, was published in Psyche, Volume 6i, p. 2 t , t 954. BACK VOLUMES OF PSYCHE The Cambridge Entomological Club is able to offer for sale the following volumes of Psyche. Volumes 3, 4, 5, 6, 7, 8, each covering a period of three years. $8.00 each. Volumes 10, 14, 17 to 26, each covering a single year, $2.00 each. Volumes 27 to 53, each covering a single year, $2.50. Volumes 54 to 65, each covering a single year, $3.00. Volume 66, covering a single year, $5.00. Some other volumes, lacking certain issues, are also available (information upon request). Orders for 10 or more volumes subject to a discount of 10%. All orders should be addressed to F. M. Carpenter, Editor of Psyche, Biological Laboratories, Harvard University, Cambridge,, Mass. 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 38, Mass. r PSYCHE A JOURNAL OF ENTOMOLOGY Established in 1874 Vol. 67 December, i960 No. 4 CONTENTS A New Acanthoctenus (Araneae: Acanthoctenidae) from Jamaica, W. I. A. M. C flickering 81 Techniques for the Study of Ant Larvae. G. C. Wheeler and J. Wheeler 87 The Female of Lucarachne beehei Gertsch (Araneae: Symphytognathi- dae) A. M. Chickering 95 Studies on North American Carboniferous Insects. I. The Protodonata F. M. Carpenter 98 Australian Carabid Beetles IV. List of Localities, 1956-1958. P. J. Darlington , Jr Ill Author and Subject Index for Volume 67 129 CAMBRIDGE ENTOMOLOGICAL CLUB Officers for 1960-61 President H. Levi, Harvard University Vice-President N. S. Bailey, Bradford Junior College Secretary G. L. Bush, Harvard University Treasurer F. M. Carpenter, Harvard University Executive Committee A. G. Humes, Boston University J. J. T. Evans, Harvard University EDITORIAL BOARD OF PSYCHE F. M. Carpenter (Editor), Professor of Entomology , Harvard University P. J. Darlington, Jr., Head Curator of Insects , Museum of Com- parative Zoology W. L. Brown, Jr., Assistant Professor of Entomology , Cornell University ; Associate in Entomology, Museum of Comparative Zoology E. 0. Wilson, Associate Professor of Zoology, Harvard University H. Levi, Associate Curator of Arachnology , Museum of Comparative Zoology 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, $5.00 to all other subscribers. Single copies, $1.25, Checks and remittances should be addressed to Treasurer, Cambridge Ento- mological Club, Biological Laboratories, Harvard University, Cambridge, Mass. Orders for back volumes, missing numbers, notices of change of address, etc., should be sent to the Editorial Office of Psyche, Biological Laboratories, Har- vard University, Cambridge, Mass. IMPORTANT NOTICE TO CONTRIBUTORS Manuscripts intended for publication should be addressed to Professor F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. Authors contributing articles over 8 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 $6.00 per page. The actual cost of preparing cuts for all illustrations must he borne by contributors; the cost for full page plates from line drawings is ordinarily $12.00 each, and the full page half-tones, $18.00 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 September i960 Psyche (Vol. 67, no. 3) was mailed April 13, 1961. The present issue of Psyche (Volume 67, no. 4) is pub- lished with the aid of a National Science Foundation Grant (G 15817) to the Cambridge Entomological Club. The Lexington Press. Inc., Lexington, Massachusetts PSYCHE Vol. 67 December, i960 No. 4 A NEW ACANTHOCTENUS (ARANEAE: ACANTHOCTENIDAE) FROM JAMAICA, W. I.1 By Arthur M. Chickering Albion College, Albion, Michigan About twenty-five species are known in this family and all have been assigned to the Genus Acanthoctenus Keyserling, 1876. These have been reported all the way from Mexico to Argentina. So far as I have been able to learn, however, no representative has ever been recorded from the West Indies. It is, therefore, of considerable in- terest to be able to report that both sexes of what I am compelled to regard as a new species have been collected in several localities in Jamaica, W. I. A holotype male and an allotype female have been selected and are described in accord with my usual formula in this brief paper. These types are being deposited in the Museum of Com- parative Zoology at Harvard College, Cambridge, Mass. Genus Acanthoctenus Keyserling, 1876 Acanthoctenus remotus sp. nov. (Figures 1-6) Holotype male. Total length 11 mm. (somewhat exaggerated be- cause of softened and stretched condition of the specimen in the region of the pedicel). Carapace 5.59 mm. long; 4.55 mm. wide opposite second coxae where it is widest ; well rounded from opposite posterior eyes to posterior border; median longitudinal thoracic groove deep and long. Eyes. Eight in three rows, two, four, two; essentially as in the Ctenidae (Fig. 1). Viewed from above, both rows recurved; viewed Tublished with the aid of a grant from the Museum of Comparative Zool- ogy at Harvard College. 8l 82 Psyche [December from in front, anterior row strongly recurved, posterior row moder- ately recurved in order to produce the characteristic placement. Ratio of eyes AME :ALE : :PME :PLE — i4:io::22:22 (measurements taken from borders of the corneas). AME separated from one an- other by about one third of their diameter, from ALE by slightly more than their diameter. PME separated from one another by about one third of their diameter, from PLE by a little less than their diameter. LE separated from one another by about twice the diameter of ALE (the corneas of these eyes are extended much fur- ther than usual). Central ocular quadrangle wider behind than in front in ratio of 10:7, wider behind than long in ratio of 25:22. AME separated from PME by a little more than one third of their diameter. Height of clypeus equal to about the diameter of AME. I he clypeus bears 17-18 long slender spines with one isolated on each side dorsal to the base of the cheliceral boss. Chelicerae. Nearly perpendicular, slightly divergent, moderately robust. Basal segment 2.275 mm. long. With many long slender spinules extending forward from promargin and many long, curved, light colored hairs at distal end. Basal boss well developed. Fang moderately robust, fairly evenly curved; with three promarginal teeth, the middle one the largest; with three retromarginal teeth of moderate size and gradually diminishing in size from near base of fang distally. Maxillae. Nearly parallel, moderately robust, only slightly concave in distal half along lateral border. Only slightly more than twice as long as broad. With well developed scopula along medial border to distal tip. Serrula only moderately well developed along distal lateral corner. With many long slender and stiff bristles especially along lateral border. Lip. Only a little longer than wide in middle; slightly concave along distal border; reaches only slightly beyond middle of maxillae. Sternal suture slightly procurved. Sternum. Subrotund; as wide as long; slightly pointed at posterior end which is not extended between fourth coxae which are separated by about one fifth of their width. Only slightly scalloped opposite the coxae. Moderately well supplied with slender stiff bristles. Legs. 1423. Width of the first patella at “knee” .845 mm., tibial index of first leg 6. Width of fourth patella at “knee” .780 mm., tibial index of fourth leg 8. 1960] Chickering — Acanthoctenus 83 Femora Patellae Tibiae Metatarsi Tarsi Totals (All measurements in millimeters) /. 8.450 2-795 10.400 9.425 3.250 34-320 2. 7475 2.600 7.280 7.930 2-795 28.080 1 A* 5.590 1.820 4.940 6.630 2.405 21.385 4- 8.190 2.275 7.150 9.230 3.315 30.160 Palp 3.120 1.300 1.625 2.015 8.160 All trochanters slightly notched ventrally at distal border. Tarsi with two claws, each pectinate in a single row. Trichobothria ob- served on femora, tibiae, metatarsi, tarsi and on palps. The distribu- tion appears to be irregular, but this has not been carefully enough studied to warrant a precise statement. Figures 1-6. External Anatomy of Acanthoctenus rcmotus. Fig. 1. Eye group of male; seen from in front. Fig. 2. First tibia; ventral view. Fig. 3. Left palpal tibia; nearly dorsal view. Fig. 4. Left palpal tibial apophysis; lateral view. Fig. 5. Left palpal tibia and tarsus; nearly ventral view. Fig. 6. Ventral view of epigynum. 84 Psyche [December Spines. There are many prominent spines on the legs but those which seem to be the most distinctive are : First tibia with seven pairs of long, fairly robust, ventral spines (Fig. 2) ; first metatarsus with ventral spines 0-2-2-2-lr-lr-lp. Second tibia with ventral spines essen- tially as in first but with more irregularity; second metatarsus with spines essentially as in first but also with more irregularity. Third tibia with three pairs of these robust spines; third metatarsus with ventral spines 02-2-2. Fourth tibia with ventral spines 2-2-2; fourth metatarsus with four pairs of ventral spines very irregularly placed. Prolateral and retrolateral robust spines easily confused with ventral spines. Calamistrum apparently absent. Tarsal scopulae well de- veloped on tarsi one and two but less well developed on tarsi three and four. Metatarsal scopulae moderately well developed on meta- tarsi one and two but absent from three and four. All tarsi with well developed claw tufts. Palp. Relatively simple; closely similar to that of A. spinigerus Kerserling from Central America. Details best shown in Figures 3-5. Tibia with robust distal retrolateral apophysis ; with two long slender dorsal spines near proximal end and a similar prolateral spine near proximal end. Patella with dorsal spines as follows: 2-0-1 (extremely long and slender). Hairs on tibia extremely long and stiff. Abdomen. Somewhat ovate with widest part about one third of its length from posterior end; 5.85 mm. long and 3.575 mm. wide at widest part. With a pair of dorsal depressions a little behind the middle. With three pairs of spinnerets, all short but with anterior pair the largest and the middle pair the smallest. With cribellum obscure and poorly developed. Color in alcohol. Carapace with a broad, median, nearly uniformly light stripe. On each side of the posterior half of the median thoracic groove there is an elongated dark spot. The whole area lateral to the median light stripe is brownish. Between the PLE there are four narrow stripes formed by light grayish hairs. The chelicerae are irregularly striped with brownish ; in front at the outer proximal corner there is a narrow stripe of grayish hairs. Abdomen : the dor- sum is brownish, much lighter in the middle in the dorsal half. There is also a group of four light colored tufts of long hairs in the posterior quarter. The whole ventral area, sternum, and venter is uniformly yellowish. The legs are generally brownish dorsally and yellowish ventrally. 1960] Chick e ring — A canthoctenus 85 female allotype. Total length 15.08 mm., including the somewhat swollen bases of chelicerae; without the latter 14.625 mm. long. Carapace 6.1 1 mm. long, 5.20 mm. wide at interval between second and third coxae where it is widest; only slightly sloping from between PLE to about the middle of the well developed longitudinal median thoracic groove from which position it descends steeply to posterior border. Eyes. Arrangement as in male. Ratio of eyes AME :ALE : :PME : PLE = 14:12 ::20:23 (measurements taken from borders of corneal hypodermis). AME separated from one another by about five sev- enths of their diameter, from ALE by about 9/7 of their diameter. PME separated from one another by about their radius, from PLE by about their diameter. Laterals separated from one another by about the long diameter of ALE. AME separated from PME by the radius of the former. Central ocular quadrangle wider behind than in front in ratio of about 5 :3-5 ; wider behind than long in ratio of about 5:4.5. Height of clypeus equal to about 17/14 of the di- ameter of AME. Clypeus with about 12 long slender spines. Chelicerae. Nearly perpendicular; parallel, robust, moderately gib- bous in front; basal segment 2.6 mm. long. Basal boss well developed. With many stiff bristles in front. Fang and teeth along fang groove essentially as in male. Maxillae , Lip , and Sternum. Essentially as in male. Legs. 1423. Width of first patella at “knee” .975 mm., tibial index of first leg 9. Width of fourth patella at “knee” .910 mm., tibial index of fourth leg 11. Femora Patellae Tibiae Metatarsi Tarsi Totals (All measurements in millimeters) I. 6.500 3.120 7.410 6.305 2.210 25-545 2. 6.125 2.795 6.1 10 5-655 1.820 22.505 3. 5-525 2.275 4-485 5.200 I.885 19.370 4. 6.825 2.275 5.850 8.190 2.340 25.480 Trichobothria observed on many segments of legs and palps but their exact distribution has not been determined. Calamistrum short and near proximal end of fourth metatarsus. Spines. First leg: femur dorsal O-i-i-i, prolateral 0-1-1-0, retro- lateral O-i-i-O-i-i-i-O, ventral with numerous spines irregularly placed and difficult to describe significantly; first tibia with nine pairs 86 Psyche [December of ventral spines; first metatarsus with five pairs of what may be termed ventral spines but most of them are moved into: a somewhat lateral position. The second tibia has seven pairs of spines and the second metatarsus five pairs, all similar to those on first leg. All legs are provided with numerous stout spines. The palpal claw is simple with four moderately robust teeth. Abdomen. Regularly ovate; cribellum moderately well developed; clearly divided. Otherwise essentially as in male. Epigynum. Essentials as shown in Figure 6. Color in alcohol. Essentially as in male except as herein noted. Carapace more clearly striped than in male; on each side of the median light stripe there is a broad darker stripe and on each an additional light marginal stripe thus making five broad stripes on the carapace. Type locality. Holotype male from St. Andrew, Upper Mountain View, January 15, 1950 (C. B. Lewis). Allotype female from St. Andrew, Cross Roads, December 3, 1950 (R. P. Bengry). The fol- lowing specimens are also in the collection : One paratype male from St. Andrew, Upper Mountain View, May 15, 1950 (C. B. Lewis) ; one paratype female taken with holotype male; another female from St. Andrew, Upper Mountain View, February, 1951 (C. B. Lewis) and two females from St. Andrew, Half-Way Tree, October 18, 1950 (R. P. Bengry). The author of this paper took one immature female in Hope Gardens, near Kingston, June, 1954. Selected References Bonnet, Pierre 1957. Bibliographia Araneorum. 2. Toulouse: Les Arti- sans de rimprimerie Douladoure. Keyserling, Graf E. von 1876. Ueber Amerik. Spinnenarten d. Unterord- nung. Citigradae. Verh. Zool. Bot. Ges. Wien. 26: 609-708. T. 7-8. F. 1-72. Roewer, C. F. 1954. Katalog der Araneae. Vol. 2. Bruxelles. TECHNIQUES FOR THE STUDY OF ANT LARVAE By George C. Wheeler and Jeanette Wheeler University of North Dakota, Grand Forks Our studies on ant larvae have been published in 30 separate articles scattered in eight periodicals, appearing over a period of 33 years. Several entomologists have expressed regret that the studies have not been combined into a monograph. We regret this too, but no founda- tion has been willing to publish so long a treatise with so many illus- trations and without economic or popular appeal, and no journal has been willing to accept it on the installment plan. The exigency of finance, then, has forced us to publish the body of the “monograph” piecemeal. This present article can be regarded as a part of the intro- duction. We are prompted to publish it at this time by requests for information concerning our techniques. Living Material When living larvae are available, we observe and record as much as possible of their external anatomy. If the larvae are active the changeability of form is noted. Preservation Any preservative may be used for ant larvae if only the external anatomy is to be studied. We prefer 85% ethyl alcohol. No special preparation is necessary; the larvae are simply dropped into the pre- servative. If the alcohol is too dilute or if the container has been overcrowded internal structures may be inadequately preserved, but the integument will still be usable for external anatomy. If large larvae are jostled the hairs may get broken. Naturally if one plans to investigate also internal anatomy or histology, appropriate killing and fixing reagents are necessary. If we have only one larva (or very few) of a species we make notes and drawings before subjecting it to any additional techniques. This is simply good insurance. Relaxation Even dried larvae may be used for the study of external anatomy. Either of two methods of relaxing is recommended : 87 88 Psyche [December 1. Two days in a 0.5% solution of trisodium phosphate. Transfer to 85% alcohol. 2. Two days (or less) in the following — 95% ethyl alcohol 280 ml; distilled water 230 ml; benzol 35 ml; ethyl acetate 95 ml. Dried larvae are handled with extreme care, since hairs are more likely to get broken in this condition. Handling The most convenient receptacle for the treatment of larvae is a staining dish ( = embryological cup) 41x41x18 mm in outside dimensions. Minute specimens, however, are best treated in culture slides (= hanging-drop slides). Small specimens are transferred with a pipette, larger specimens between the points of forceps (but without compression). Or, if one prefers, the larvae may be left in the same dish or culture slide; the old reagent is drawn off with a pipette and then replaced with the next reagent. Cleaning The best cleaning reagent is potassium hydroxide solution (10 gm KOH in 90 ml water). While still in preservative the larvae are punctured with a dissecting needle or minute insect pin on the right side to permit the ready penetration of the cleaning solution. The number and locations of the punctures and the size of the needle de- pend upon the size of the larva. The specimen is left in the cleaning solution until all the internal tissues are dissolved and only a trans- parent exoskeleton remains. If cleaning is not complete by the end of two days, the following procedure may prove effective: transfer to 1% hydrochloric acid and leave 15 minutes; 15 minutes in 95% alcohol; then leave in KOH until clean. Some larvae contain droplets of opaque substances which are insoluble in KOH ; these usually disappear later in alcohol or xylol. Staining The exoskeletons of ant larvae are stained in a very dilute solution of acid fuchsin. We use the following formula as a stock solution: acid fuchsin 0.1 gm, concentrated hydrochloric acid 1 ml, distilled water 1 liter ; a few thymol crystals are added to prevent mold. The cleaned exoskeletons are washed in water for 15 minutes and trans- ferred to 2 ml of 1% HCL in a staining dish; five drops of the stock solution of acid fuchsin are added. The integuments are left in the stain for 12 hours. We have experimented with stronger solutions of stain for shorter periods but have not been satisfied with the results. 1960] Wheeler — Ant Larvae If overstained the integument can be completely decolorized by leaving it in a i% solution of potassium hydroxide for several hours. It can then be washed and restained. Dehydration After staining, the exoskeletons are transferred to 95% alcohol (about 3 ml), in which they remain for about an hour. At the end of this period eight drops of carbolxylol ( = 3 parts xylol and 1 part melted phenol crystals) are added every five or ten minutes until 48 drops have been added to the alcohol. If the carbolxylol is added too rapidly the exoskeletons may collapse and become distorted; they can be restored by transfer to 95% alcohol. If oil droplets separate from the alcohol, too much water has been carried over from the stain ; the integuments are transferred to fresh 95% alcohol. Clearing From the mixture of carbolxylol and alcohol the integuments are transferred to pure carbolxylol and left for 15 to 60 minutes; then to xylol in a culture slide (or if large, in a staining dish). Add a drop of thin Canada balsam every 15 minutes, allowing the xylol to evapor- ate meanwhile. Mounting When the balsam in the culture slide attains the same consistency as that which is being added, a large drop of the latter is placed on a slide and the exoskeleton is transferred to it. At this time medium to large specimens are surrounded — or at least flanked — by supports of some sort. We use fragments of broken slides which are of about the same thickness as the depth of the specimen. If it is obvious that the exoskeleton will not remain in the desired position, supports (such as a fine glass rod or fragments of cover slip) may be placed under the low part. Positioning With needles dipped in xylol the integument is now arranged in the proper position for drawing (discussed below). Since the balsam must be soft at this stage, the specimen will rarely remain in precisely the desired position ; hence the slide is allowed to rest flat in a covered con- tainer (e.g., a petri dish) for 24 hours. If the specimen then needs rearranging, this is done under a stereo- microscope with needles dipped in xylol. If the balsam is too firm a drop of xylol is put on it and let stand a few minutes before using needles. If it is apparent that the specimen will not remain in the 90 Psyche [December desired position, supports (fine glass rods or fragments of cover slip) may be inserted at this time. If supports were placed earlier, they may need readjustment. Shifting the retaining walls will sometimes correct the position of a specimen. After the rearrangement is completed more balsam is added if necessary to cover the specimen. It is sometimes necessary to rearrange a specimen on half a dozen successive days. Once a specimen has remained in the desired position for 48 hours and is completely covered with balsam, the technique is considered finished. If material is abundant we regularly make two kinds of mounts of each size and caste of each species: (1) an entire exoskeleton in side view; (2) detached head in full-face view, with the remainder of the exoskeleton ventral side up (or, if the head is applied to the ventral surface, the whole exoskeleton ventral side up). Other kinds of mounts are made whenever necessary. For a side view an entire exoskeleton with the left side up and in strict profile, i.e., the middorsal line, the midventral line, the middle of the anus and the middle of the labium are all in the same plane. A good check is to be sure that the left spiracle of each somite is directly above the right spiracle of the same somite. It is conventional to show the entire larva viewed from the left side. Actually it doesn’t matter which side is up. If for any reason (e.g., damaged left side) an in- tegument is mounted with the right side up, it is a simple matter to reverse the original drawing. The exoskeleton of the head is mounted in full-face (i.e., anterior) view. To accomplish this the uppermost line (imaginary) of the oc- cipital border is set in the same plane as the lowermost part of the ventral surface of the labium. When the head is applied to the ventral surface of the body (i.e., no neck), it is often possible to get it in the correct position by mounting an entire integument ventral side up. In most genera, however, it is necessary to cut off the head (either alone or with a portion of the prothorax) and mount it separately. It is often necessary to prop up the ventral part of the head on fragments of cover slip, but only very large heads require retaining walls. After the head is removed, the remainder of the integument is mounted with the ventral side up by following the procedure for the entire integument (see above). 1960] Wheeler — Ant Larvae 91 Covering It is not, however, necessary to cover a mount if it is to be studied at low magnifications. In fact, covering is avoided whenever possible; if the balsam is soft (and it may take years for the interior of a thick mount to harden), the weight of the cover slip will set up currents which are likely to move the specimen out of the desired position. If, however, the specimen must be studied under the high power objective (X440) of a microscope, a cover slip is essential. For study under an oil immersion objective (X970) the part of the integument is detached, flattened and covered. If a cover slip is required on a thick mount, it may be added at any time after the surface of the balsam is hard. A drop of xylol is put on the surface and a large drop of thin balsam is put on a cover slip. The cover slip is inverted and one edge is rested on the balsam mount; the opposite edge is supported by a dissecting needle while it is slowly lowered into position. The cover slip is then adjusted by needles dipped in xylol. If material is scarce — and especially if there is only a single larva — low-power studies are completed before the mount is covered. When material is abundant both covered and uncovered mounts are prepared. Reversibility It is reassuring to realize that, after the integument has been cleaned, every step is reversible. The same may be said of any series of steps and even of the entire technique from cleaned exoskeleton to covered slide mount. Only one precaution is necessary: hydration (i.e., from 95% alcohol to stain) must be gradual. Perfectionism The most important advice we can give to beginners is to avoid perfectionism in making mounts of ant larvae. Not only is perfection- ism harmful to the technician (frustration, waste of time), but it can be very damaging to the material. Whenever the conscientious worker has an almost-good-enough preparation, there is always the tempta- tion to try to adjust it a little bit more. Since the balsam must be a little thicker than usual to hold the specimen in the new position, the risk of damage is therefore proportionately increased and the damage may well be irreparable. It is far better to stop short of perfection in mounting and resort to correction of the position of the slide. Before drawing we make 92 Psyche [December any slight correction that may be necessary by propping up an edge or a corner or an end of the slide on a small lump of plasticine. Storage Prepared slides of ant larvae are always stored flat in the horizontal position. To store on edge is to risk displacement of the specimen, since the interior of a thick balsam mount requires years to harden. Thin mounts may, of course, be stored in any position as soon as the edges of the cover slip are sealed. Preserved larvae may be stored indefinitely in 85% alcohol. The only precaution required is to prevent drying by evaporation through a faulty seal. (Dried larvae may be relaxed by methods given above.) Summary of T cchnique 1. Puncture integument 2. 10% KOH until clean 3. Distilled water — 15 minutes 4. Transfer to 1% HCL 5. Add 5 drops acid fuchsin ; leave 12 hours 6. 95% alcohol for an hour 7. Add carbolxylol gradually 8. Carbolxylol 15-60 minutes 9. Xylol 10. Add thin balsam gradually 11. Transfer to balsam on slide 12. Arrange in desired position 13. Examine daily; rearrange as of- ten as necessary until desired po- sition has been maintained for +8 hours Descriptions We have generally followed this plan in writing our specific des- criptions of ant larvae : body shape ; leg vestiges ; spiracles; integument ; body hairs; head shape; antennae; head hairs; labrum; mandibles; maxillae; maxillary palp; galea; labium; labial palp; opening of sericteries ; hypopharynx. To facilitate taking notes we have pre- pared and used a mimeographed form with a blank space under each of the above headings. When a larva is to be described the prepared slides are studied under a stereomicroscope for low magnifications and a microscope for high magnifications. Measurements are made by comparison with the squares of an eye- piece reticule (= net micrometer) which has been calibrated with a stage micrometer. D r azvings All our drawings of ant larvae are made with the aid of a micro- scope. A reticule (= net micrometer) is placed in the eyepiece. This superimposes a grid of squares on the image of the object. The draw- ing of the object is made on coordinate paper. The size of squares used is determined by the desired enlargement of the drawing. If 1960] Wheeler — Ant Larvae 93 coordinate paper is not available or if the squares are not of the desired size, it is not difficult to construct a coordinate grid on plain paper. When the eyepiece grid is properly placed with reference to the image and the coordinate system is established on the paper, we are ready to start drawing. The dimensions and proportions of the drawing are established by short marks across the lines of the coordinate system in the appropri- ate places. The main outline is constructed by joining these inter- sections with freehand lines. Structures inside or outside the main outline are treated in approximately the same way, but more freehand drawing is required because fewer squares are involved and smaller parts are likely to be wholly inside a single square. It is not necessary to complete all fine details at this stage. The drawing is placed on a tracing table, a sheet of thin drawing paper (e.g., Ledger Linen) is laid on it; the drawing is copied onto the drawing paper in India ink with curve-pen (Keuffel and Esser). Finer details are drawn with a Hunt Mapping Pen No. 104. In the case of bilaterally symmetrical structures (e.g., head in anterior view) only one half is drawn, with a vertical matching line down the middle. After this half (but not the matching line) has been inked the pencil drawing is turned over on the tracing table, matched up to the first half and then inked. Most of our drawings are orthographic projections of opaque ob- jects (even though the integuments studied are actually transparent). Only structures on the near side of the plane of the outline are drawn. For example, in a side view of a larva the outline represents the imaginary middorsal and midventral lines; hairs and other parts attached beyond (= below, in the field of the microscope) the plane of these two lines are not shown. To our rule of assumed opacity a few exceptions are made, e.g. dark-colored structures overlaid by translucent tissues. We have found 120 mm to be a convenient length for a drawing of the entire larva. The plane of projection includes the middorsal line (imaginary), the midventral line (imaginary), the middle of the anus and the middle of the bottom of the labium. It is convention- al to view the larva from the left. But if for any reason it has been drawn from the right, the original drawing (on co-ordinate paper) can be turned over on the tracing table before inking. The head is drawn in full-face (i.e., anterior) view. The plane of projection passes through the uppermost line (imaginary) of the 94 Psyche [December occipital border and the lowermost part of the ventral surface of the labium. A convenient width (at the widest place) is 75 mm. The mandible is also drawn in anterior view. The plane of pro- jection passes through the apex, the inner (= medial) condyle and the outer ( = lateral) condyle. If the mandible is drawn from a head mounted for full-face view, it is usually necessary to prop up an edge of the slide to bring these three points into the same horizon- tal plane. A convenient length for the mandible drawing is 65 mm. THE FEMALE OF LUCARACHNE BEEBEI GERTSCH (ARANEAE: SYMPHYTOGNATHIDAE) 1 By Arthur M. Chickering Albion College, Albion, Michigan Miss Elizabeth B. Bryant established the genus Lucarachne in 1940 on the basis of both males and females taken in Oriente Province, Cuba, at 1500 feet elevation. She placed the genus, with some hesita- tion, among the Metinae of the Argiopidae and noted that it appeared to be closely related to Theridion cidrelicola Simon from Venezuela, the male of which is now considered to belong to the genus Lucarachne and is included in the Family Symphytognathidae. When Dr. Gertsch (i960) described L. beebei from males alone he noted that four spe- cies of the genus Lucarachne are now known as follows: L. tibialis Bryant from Cuba; L. cidrelicola (Simon) from Venezuela; L. beebei Gertsch from Trinidad; L. palpalis Kraus from Honduras and Mexico. Dr. Gertsch has recently identified a male L. beebei Gertsch from my Panamanian collection of 1936 taken in El Valle. In my collection made in Panama in 1958 numerous specimens of both males and females of this last-named species were found also near El Valle. One of the females has been selected and is described below in accord with my usual procedure. Genus Lucarachne Bryant, 1940 Lucarachne beebei Gertsch Female. Total length 2.05 mm. Carapace 1.012 mm. long, .85 mm. wide opposite interval between second and third coxae where it is widest ; well rounded along margin from opposite posterior eyes to posterior border ; gently arched from PE to near posterior border with highest part shortly behind those eyes ; with few bristles or hairs. Eyes . Eight in two rows. Region of AME considerably raised into a low blunt tubercle. Seen from above, posterior row nearly straight, anterior row strongly recurved ; seen from in front, anterior row slightly procurved, posterior row strongly so, all measured by centers. Tublished with the aid of a grant from the Museum of Comparative Zool- ogy at Harvard College. 95 96 Psyche [December Ratio of eyes AME :ALE : :PME :PLE = 10:61:6:7. AME separ- ated from one another by about three fifths of their diameter, from ALE by only a line. PME separated from one another by about five sixths of their diameter, from PLE by about the same distance. LE almost in contact. Central ocular quadrangle wider in front than behind in ratio of about 6 :5 ; nearly as long as wide in front. Clypeus very high; height equal to nearly three times the diameter of AME. Figures 1-4. External Anatomy of the Female of Lucarachne beebei Gertsch. Fig. 1. Left first femur; nearly ventral view. Fig. 2. Eye group of female; somewhat obliquely from above. Fig. 3. Epigynum, from below. Fig. 4. Abdomen; posterior surface. Chelicerae. Moderately robust, vertical, parallel; fang regularly curved ; promargin of fang groove apparently with three teeth the first of which is only a denticle while the other two are fairly robust; retromargin with only one rather small tooth. Maxillae. Robust ; slightly convergent. Lip. Reaches about two thirds the length of the maxillae. Sternal suture only slightly procurved. Sternum. Scutiform in general; quite convex; only slightly longer than wide; widest opposite interval between first and second coxae; extends between fourth coxae and terminates in a blunt truncature; fourth coxae separated by nine tenths of their width ; posterior end with a cluster of stiff bristles. Legs. 1243. Width of first patella at “knee” .132 mm., tibial index of first leg 9. Width of fourth patella at “knee” .1 10 mm., tibial index of fourth leg 10. 1960] Chickering — Lucarachne beebei 97 Femora Patellae T ibiae Metatarsi T arsi Totals (All measurements in millimeters) I. 1-275 .422 1. 000 .715 .620 4-032 2. 1. 150 -390 .785 .715 .520 3.56o 3- .910 .325 ■585 .484 .484 2.788 4 - .970 -340 •715 .715 .585 3.325 Legs with many long slender stiff bristles and few definite long slender spines the most significant of which appear to be the following: each patella with two of these dorsally placed, one near each end of the segment; each tibia has one of these on the dorsal surface near the proximal end ; numerous extremely slender spines occur more or less in rows on the ventral surfaces of the femora. The first femur has a distinctive ventral apophysis near the distal end (Fig. i). Tricho- bothria are present on the tibiae and metatarsi but the exact numbers and distribution have not been determined. Abdomen. Ovoid; carried in nearly a perpendicular position; over- laps the carapace considerably; similar to that of L. palpalis Kraus (Fig. 4) . Epigynum. Simple; details best shown in Figure 3. Color in alcohol. Essentially as described for the male by the author of the species; exposed posterodorsal surface essentially as shown in Figure 4. Collection records. Dr. Gertsch had the male holotype and one male paratype from Trinidad. Collection records in Panama are as follows: a single male from El Valle from my collection of July, 1936, recently identified by Dr. Gertsch; several specimens of both sexes are included in my collection of 1958; these were taken about five miles south of El Valle in January. References Cited Bryant, Elizabeth, B. 1940. Cuban Spiders in the Museum of Compara- tive Zoology Bull. Mus. Comp. Zool. at Harvard College, 86 (7): 249-532, 22 pis. Forster, Raymond R. 1958. Spiders of the Family Symphytognathidae from North and South America. American Museum Novitates, No. 1885:1-14. Gertsch, Willis J. 1960. Descriptions of American Spiders of the Family Symphytognathidae. American Museum Novitates, No. 1981. Kraus, Otto 1955. Spinnen aus El Salvador ( Arachnoidea, Araneae). Abhand. der Senckenbergischen Naturforschenden Gesellschaft. 493: 1-112. 12 tafeln. STUDIES ON NORTH AMERICAN CARBONIFEROUS INSECTS 1. THE PROTODONATA1 By F. M. Carpenter Harvard University Introduction This is the first of a series of papers based upon insects from Car- boniferous strata in North America. The studies will be concerned in the main with previously described specimens (mostly types) in the Museum of Comparative Zoology, the U. S. National Museum and the Peabody Museum at Yale University, but will include new material whenever it is available. Order Protodonata Up to the present time only three species of Protodonata have been described from North American deposits of Carboniferous age. The present paper includes the description of a fourth species and an account of the previously described ones. In addition, I have in- cluded a discussion of the status of the ordinal name Protodonata. One of these protodonates belongs to the family Meganeuridae and another to the Paralogidae. The remaining two are best referred to Incertae Sedis for family designation, although their protodonate affinities are unquestionable. Family Meganeuridae Typus durhami, n. sp. Plate ii ; text figure i. Forewing: greatest length of preserved part, 88 mm.; estimated complete length, 175 mm.; maximum width of preserved part, 27 mm. The venational details of both fore and hind wings, so far as preserved, are shown in text figure 1. The general venational pattern is close to that of permianus and other known species of the genus. The cells of the wings are larger than those of permianus , especially in the area between MA and CuP; R2 + 3 and R4 + 5 seem to diverge even more gradually in durhami than in permianus. The basal origin of Hhis research was supported by National Science Foundation Grant No. NSF G 14099. and the article is published with the aid of a grant from the Museum of Comparative Zoology at Harvard College. 98 1960] Carpenter — Protodonata 99 R 3 is not preserved in the fore wing but in the hind wing it is clearly visible and is associated with the oblique cross-vein (Sn) characteristic of most Meganeuridae (Carpenter, 1947, p. 47) \ the cross-vein is not so pronounced as it is in permianus, however. 1 he differences between the fore and hind wings in the proximal region are like those in other species of Typus. Holotype: U. S. National Museum, Washington, D. C.2 The specimen (field number 8758) was collected in May, 1939, by Mr. Charles B. Read at Durham, Georgia, the locality data being “Catoosa (TVA. Durham Quad.), Williams Coal Company Mine in No. 4 Coal. Plants from roof shale.” According to Butts’ account of the stratigraphy of the Paleozoic area in northwest Georgia (Butts and Gildersleeve, 1948, pp. 54-56), the coal mines at Durham are in the Walden sandstone part of the Pottsville Series. Butts states that “according to the best knowledge, the Pottsville Series of Georgia is of Lower Pottsville age and falls within the limits of the Lee Con- glomerate of Tennessee and Virginia and corresponds approximately to the lower part of the Pottsville of the anthracite coal fields of Pennsylvania which carry the Lykens number 4 and 5 and the Lykens Valley coals.” A brief discussion of the nature of the no. 4 coal at Durham is contained in Gildersleeve’s account (ibid., 1948, p. 104) of the mineral resources of the Paleozoic area in northwest Georgia. As can be seen from the photograph ( Plate 1 1 ) , the specimen con- sists of the apical portions of the fore and hind wings; although the remainder of the insect, which was presumably originally preserved, is broken away, the preserved portions are very clear. I have assigned this species to the genus Typus but it might conceivably belong to Megatypus or Boltonites ; the absence of the basal part of the wing containing the regions of the anal crossing and the second anal vein prevents a more definite generic assignment. The interest attached to this species is in the evidence which it pro- vides for the existence of the Meganeuridae in North America during the Carboniferous. It is the first such fossil that can be definitely assigned to the family. Furthermore, Typus durhami, occurring in the Lower Pottsville Series (corresponding approximately to West- phalian A), is among the oldest fossil insects known from North America. ‘I am indebted to Dr. S. H. Mamay of the United States Geological Survey for sending me this specimen for study. Psyche, 1960 Vol. 67, Plate 11 Carpenter — Protodonata 1960] Carpenter — Protodonata IOI Family Paralogidae 1 he distinctive features of this family are the short subcosta, which terminates just beyond the middle of the wing; and the widely di- vergent branches of Rs (R2 + 3 and R4 + 5). The complete absence of the apparent vestige of CuA may also be a family trait. Paralogus is the only Carboniferous genus that clearly belongs here but in my opinion the Permian genus Oligotypus Carp, does also. Fraser (1957) considered Oligotypus to be a meganeurid, but, as shown below, his concept of the Paralogidae has not been entirely correct. Oligotypus Text figure 1. Drawing of Typus durhami , n. sp. (holotype), fore and hind wings. Sc, subcosta; Rs, radiust; R2, R3, Rl+5, branches of radial sector; MA, anterior media; CuP, posterior cubitus; Sn, subnodal vein. differs from Paralogus only in minor details: the wing is somewhat more slender, and the branches of MA and iA arise as a series of irregular veinlets, instead of distinct branches as in Paralogus. The following review of the structure of Paralogus strengthens my con- viction that it and Oligotypus belong to the same family. Explanation of Plate 11 Photograph of Typus durhami, n. sp. (holotype) X 11/10. 102 Psyche [December Paralogus aeschnoides Scudder Text figure 2. Paralogus aeschnoides Scudder, 1893, U. S. Geol. Surv., Bull. 101: 21; pi. 1, fig. a. The unique specimen on which this species was based was collected by Mr. F. P. Gorham ( 1889) in Upper Carboniferous rocks at Silver Spring, East Providence, Rhode Island, and was donated to the Museum of Comparative Zoology by Professor Gorham in 1932. It consists of a well-preserved wing, about two-thirds complete. Scudder’s drawing of the fossil is good and even shows by means of shading the convexity and concavity of the veins. I have included here a new description and an original illustration of the fossil, in part because a few details of phylogenetic significance were not clearly indicated by Scudder and in part because Dr. Fraser’s recent illustra- tion of the fossil (presumably based on Scudder’s drawing) is mis- leading in several important respects. Text figure 2. Drawing of Paralogus aeschnoides Scudder (holotype). Lettering as in text figure 1. The wing fragment, as preserved, is 54 mm. long, and has a maxi- mum width of 19 mm. ; the complete wing was probably about 80 mm. long. T he subcosta terminates a short distance beyond the middle of the wing and beyond the point of separation of R2 + 3 and R4 + 5. The two latter veins diverge widely after their origin ; MA has a series of pectinate branches beginning just beyond the divergence of R2 + 3 and R4 + 5; just before the level of this divergence, iA sepa- rates into a divergent fork and sends a series of additional branches towards the distal part of the wing. The wing itself is broad for a protodonate and has a strongly curved posterior margin. However, on the basis of our knowledge of the differences between the fore and hind wings of Meganeuridae, I 1960] Carpenter — Protodonata 103 believe that the specimen of aeschnoides is a fore wing. At any rate, the distance between the posterior margin of the wing and the first anal is like that of other protodonate fore wings, not hind wings. As can be seen from Scudder’s illustration, the wing was subjected to some distortion in the process of preservation; it rests on a very uneven surface of the rock, so that the contour of the anterior margin of the wing is difficult to follow exactly. This is made worse by the presence of a slight “fault” extending obliquely across the wing, so that the veins in the anterior part of the wing are not quite aligned on the two sides of the fault. The result of the faulting and of the irregularity of the surface of the rock is to cause a more pronounced curvature of the anterior margin of the wing than would presumably otherwise have been present. The drawing in text figure 2 has been made without any attempt to restore the presumed original shape of the wing, apart from aligning the veins across the fault. Scudder’s representation of the shape of the wing is correct, the pro- portions of his drawing being approximately the same as those in the fossil. On the other hand, Fraser’s drawing (1957, figure 11), show- ing a markedly broad wing and strongly curved posterior margin, is apparently incorrect. At any rate, I do not know of any evidence which supports this conception of the wing. Two other, more minor, corrections in Fraser’s figure should be noted. The vein which he has labelled R3 is a convex intercalary vein ; R2 and R3 presumably sep- arate much further along the wing. Also, the subcosta terminates gradually somewhat beyond the level of the separation of R2 + 3 and R4 + 5, not abruptly before this level as shown in Fraser’s drawing. The short basal vein, termed CuA, which is consistently present in the meganeurids, is not discernible in the specimen of aeschnoides although Dr. Fraser has shown it in his drawing. "The absence of this vein appears to be another characteristic shared by Paralogus and Oligotypus. The deposit in which the specimen of aeschnoides was found is usually referred to the Allegheny or Conemaugh Series, about equiva- lent to the Upper Westphalian of Europe. Family: Incertae Sedis Paralogopsis longipes Handl. Text figure 3. Paralogopsis longipes Handl., 1911, Amer. Journ. Sci., (4) 31: 374, fig. 58. The specimen on which this species was based is contained in an ironstone nodule from the vicinity of Mazon Creek, Illinois; the type 104 Psyche [December is in the Peabody Museum at Yale University, where I examined it several years ago. Although Handlirsch figured only the hind wing (so far as it was preserved), a portion of the fore wing is also present; this is clearly narrow, more like that of the Meganeuridae than of the Paralogidae, where Paralogopsis has previously been placed (Fraser, 1957). However, since the key parts of the wings (such as the fork- ing of Rs) are not included in the preserved portion, the family rela- tionships remain obscure. Text figure 3. Drawing of Paralogopsis longipes Handl. (holotype). Let- tering as in text figure 1. The significance of this fossil is that it provides the only record of the Protodonata in the Carboniferous strata of the Eastern Interior Region of North America. The Frances Creek shales, which yield these ironstone nodules, are considered part of the Carbondale Forma- tion ; this is regarded as about equivalent to the middle or late W est- phalian stage (late C or early D) of Europe. 1960] Carpenter — Protodonata 105 Palaeotherates pennsylvanicus Handl. Text figure 4. Palaeotherates pennsylvanicus Handlirsch, 1906, Proc. U. S. Nat. Mus., 29: 690, fig. 17. Palaeotherates pensilvanicus [sic] Handlirsch, 1906, Fossilen Insekten: 311; pi. 32, fig. 5. The fossil on which this species was established consists of a wing fragment preserved in black shale; it was collected in 1887 in inter- conglomerates, at Coxton, one mile north of Pittston, Pennsylvania. The type specimen, which is in the U. S. National Museum (No. 38787), was kindly loaned to me for study through the courtesy of Dr. G. A. Cooper. Handlirsch’s drawing of the fossil, although correctly representing the general venational features, omitted two significant details, — the subnodal vein and the costa. He recognized that the “second vein” must be the radius (Ri), but since he could detect no anterior vein, other than a marginal one, he concluded that the subcosta had fused with the costa in the area of the wing preserved. However, the clear preservation of the subnodus (S11), which Handlirsch did not figure, shows that the part of the wing represented was too near the middle of the wing for the termination of the subcosta to have taken place. Furthermore, careful study of the fossil shows that the anterior margin of the wing (costa) is actually present as distinct from the subcosta in the distal part of the fossil, although it is broken away along the rest of the wing fragment. It now becomes clear that the fossil represents a fragment of the wing just beyond the middle; it includes the point of separation of R2 and R3, but not the separation of R2 + 3 and R4+5. Handlirsch’s naming of the veins is incorrect; the convexities and concavities, which are clearly preserved in the fossil, show that R4 + 5 was included in the complex which he termed the media. The original insect was probably about the size of most species of Typus , not “very large” as estimated by Handlirsch. The wing frag- ment is 45 mm. long, and on the basis of comparisons with other protodonates, it probably represents about one-third of the complete wing. Since the width of the wing of pennsylvanicus is 18 mm., its original dimensions were probably close to those of Typus gracilis Carp. (Permian), which is 145 mm. long and 26 mm. wide. The drawing of the fossil shown in text figure 4 is based on the type specimen. The venational pattern, so far as it is known, is like io6 Psyche [December that of the meganeurids, but, since the proximal half of the wing is unknown, I consider the family position uncertain. It is clearly not a member of the Paralogidae, where it was placed by Fraser (1957). The deposit in which this fossil was collected appears to be of upper Pottsville age corresponding to Westphalian C of the European classi- fication. Status of the Ordinal Name Protodonata I take this opportunity to discuss the status of the ordinal name Protodonata, in an attempt to clear up some of the confusion which has appeared in the literature in recent years. Handlirsch (1906b) first defined this group as an order in 19063 and he placed here three new Text figure 4. Drawing of Palaeotheratcs pennsylvanicus Handl. (holo- type). Lettering as in text figure 1. 3Much confusion has resulted from two, almost simultaneously published works by Handlirsch, both appearing in 1906. His Revision of American Paleozoic Insects (Proceedings of the U. S. National Museum, Vol. 29, pp. 661- 820) is an English translation of a specially prepared account of the Paleozoic insects then known from North American deposits. New species, genera and higher taxa were described and illustrated. His Die Fossilen Insekten was published by Englemann in Leipzig in parts over a period of two years. The first part, consisting of pages 1-640, was published in 1906 (See Fossilium Catalogus , (I), Pars 16, 1922, p. 10). This part dealt with the Paleozoic insects and included, among the others, descriptions and figures of the North American fossils. That this work was published after the English account is shown by the references in Die Fossilen Insekten to specific pages in the Proceedings article. Unfortunately, many (but not all) of the species, genera and higher taxa already described in the Proceedings account were described again and designated as new in the larger, German treatise. In reality, the first designation of the Protodonata as an order was made in the Proceedings article mentioned above. The discussion of the character- istics of the order was essentially that given later in Die Fossilen Insekten , but only two genera, Paralogus and Palaeotherates , were actually mentioned as included in the order. 1960] Carpenter — Protodonata 107 families: Protagrionidae Handl., Meganeuridae Hand!., and Paralogi- dae Handl. The Protagrionidae4 were known only from a single wing (Protagrion audouini) , but the Meganeuridae were known from many specimens, some of which included the body structures. Handlirsch’s definition of the order was accordingly based on the meganeurids and the order itself was considered by him to be related to both the Paleodictyoptera and the Odonata. In the course of several years, as more meganeurids and Paleodictyoptera were found, it became in- creasingly evident that the Protagrionidae were not at all closely related to the Meganeuridae or to the Odonata. In 1932, Martynov (1932a), after pointing out the differences between these two groups, removed the Meganeuridae and Paralogidae from the Order Pro- todonata and placed them in a new order which he erected for them, the Meganisoptera. The Order Protodonata was therefore left with the family Protagrionidae, which was then recognized as having no odonate affinities. In 1943, after a detailed study of the original specimen of Protagrion audouini (preserved in the Paris Museum), I published an account of this fossil and transferred the family Protagrionidae to the Paleodic- tyoptera, where it clearly belongs, in association with several related families ; at the same time I restored the Meganeuridae and Paralogi- dae to the Order Protodonata, as they were originally placed by Handlirsch. In 1957, in h is account of the classification of the Odonata (p. 21), Fraser agreed to the separation of the Protagrionidae from the Meganeuridae and Paralogidae, but he insisted that the name Proto- donata must be associated with Protagrion , and contended that I was in error in placing the Meganeuridae in the Order Protodonata. His reason for these assertions was that “Brongniart established his family Protagriidae [Protagrionidae] on a single genus Protagrion (1885) ; the family was therefore a monotypic one and by the International Rules of Nomenclature it matters not whether the family afterwards assumed ordinal rank or that further genera or families were added to it, it must take the characters from Protagrion , that is, from the original type.” Throughout this article I am using the name Protagrionidae, instead of Protagriidae, for the family based upon the genus Protagrion. This is in accordance with the information provided by Professor Joshua Whatmough of Harvard University and published in B. E. Montgomery’s article on this subject (Annals Ent. Soc. Amer., 47: 473-474, 1954). io8 Psyche [December It is clear from this quoted passage that Fraser’s conclusions are based on his belief that Protagrion was designated by Brongniart as the type genus of the Order Protodonata and that the Rules of Nomenclature consequently require us to use that conception of the order. I believe it is Dr. Fraser who is in error here. The Rules of Nomenclature have not been applied by the Commission on Nomen- clature to orders and higher taxa. The accepted policy regarding these higher categories has been well summarized by Simpson in his recent discussion of the principles of taxonomy (1961, p. 30) : “Pro- posals have been made to extend the type system (and priority) to names of still higher taxa, above superfamilies, but this provision is not now embodied in the Rules or in general usage. At present the names of those higher taxa, of course much less numerous than names of genera or species, are determined only by consensus and acceptance of authority, and at these levels that informal system seems to work at least as well as the Rules do at lower levels.” Since the term Protodonata was first used in an ordinal capacity by Handlirsch ( 1906b), as I have stated above, and since his definition of the order was based mainly on the Meganeuridae, I prefer to use the name Protodonata for the order containing the Meganeuridae. Moreover, if the Rules of Nomenclature are applied to the ordinal name here, I contend that Protagrion has no standing as the type genus of the Protodonata and that, in fact, the genus Meganeura more logic- ally and appropriately stands as the type genus. In this connection it is necessary to correct Fraser’s statement quoted above, that “Brong- niart established his family Protagriidae on a single genus Protagrion (1885)”. At no time did Brongniart ever use the family name Pro- tagriidae [ Protagrionidae] ; it was first used by Handlirsch in 1906 (1906b). Consequently, this statement by Fraser has no meaning or application whatsoever to the term Protodonata. The name Pro- todonata was first used by Brongniart in 1885 (p. 55). In this paper Brongniart discussed a series of orders, one of these being the “Neurorthopteres” and another the “Pseudoneuroptera”. In this latter order he placed, among others, two families, one which he called Megasecopterida and another which he designated Protodonata. His precise statement about the “family Protodonata” is as follows: “Je rangerai a cote de ces Megasecopterida un type ancestral des Libellules ; la creation de la famille des Protodonata . . . et du genre Protagrion . . . me semble necessaire. Une aile seulement a ete trouvee jusqu’ici a Commentry; elles [sic] mesure 10 centimetres de 1960] Carpenter — Protodonata 109 long et 2 centimetres de large. Sa forme, sa nervation et sa reticula- tion rappellent beaucoup celle des Odonates actuels. II y a cependant d’assez notables differences.” Since this was the first mention of the genus Protagrion in the literature anywhere and since there was no mention of any species in the genus, the name Protagrion was a nomen nudum. This use of the term Protodonata did not, therefore, establish a precedent or policy with respect to use of the name. The next use of the term Protodonata, again for a family, was by Brongniart in 1894 (P* 394)- At the be- ginning of Chapter V, entitled “Protodonata”, there was a discussion of the characteristics of the Protodonata based almost entirely on Meganeura ; the genus Protagrion was not even mentioned there. In the descriptive part of the work, the first genus considered was Meganeura , which was fully described, along with two species in the genus, monyi and selysii. The other genera also assigned in the “family Protodonata” in chronological order were: Paralogus , Titan- ophasma , Protagrion , Campyloptera , and Brodia. Protagrion was, therefore, fourth in the series of genera placed in the “family Pro- todonata”. From this I think it is clear that there is no basis for Dr. Fraser’s statement that the Order Protodonata “must take the charac- ters of Protagrion Since the first definition of the Protodonata as an order (Hand- lirsch, 1906b) was based mainly on the Meganeuridae, and since Rrongniart’s first valid use (1894) of the term Protodonata for a “family” was based almost exclusively on Meganeura , I do not accept the removal of the Meganeuridae from the Protodonata and the erection of another order ( Meganisoptera) for that family. In my opinion the Order Meganisoptera is identical with the Order Pro- todonata. Whether the Protodonata should be considered a separate order or a suborder of the Odonata is largely a matter of personal choice. Fraser (1957, p. 24) considers the group (Meganisoptera) to be a suborder, whereas Martynov (1932b, p. 43; 1938, p. 62) treated it as a distinct order. I strongly support its ordinal rank; the absence of a nodus, a pterostigma and a true arculus places these insects outside the phylogenetic complex of the Odonata. no Psyche [December References Brongniart, C. 1885. Les insectes fossiles des terrains primaires. Bull. Societe Amis des Sciences Naturelles de Rouen, 1885: 50-68. 1894. Recherches pour servir a l’histoire des insectes fos- siles des temps premaires. Saint-Etienne (1893), pp. 1-493. Butts, C. and B. Gildersleeve 1948. Geology and mineral resources of the Paleozoic area in northwest Georgia. Georgia Geol. Surv., Bull. 54: 1-176. Carpenter, F. M. 1943. Studies on Carboniferous insects from Commentry, France, Part 1. Geol. Soc. Amer., Bull. 54: 527-554. 1947. Lower Permian insects from Oklahoma.. Part 1. Proc. Amer. Acad. Arts and Sci., 76: 25-54. Fraser, F. C. 1957. A reclassification of the Order Odonata. Royal Zool. Soc. New South Wales. Pp. 1-133. Handlirsch, A. 1906a. Revision of American Paleozoic insects. U. S. Nat. Mus., Proc., 29: 661-820. 1906b. Die Fossilen Insekten. Leipzig, pp. 304-311. 1911. New Paleozoic insects from the vicinity of Mazon Creek, Illinois. Amer. Journ. Sci. (4) 31: 297-377. Martynov, A. B. 1932a. New Permian Palaeoptera with the discussion of some problems of their evolution. Trav. Inst. Paleozool. Acad. Sci. 1: 1-44. 1932b. On the wing-venation in the family Meganeuridae (Meganisoptera) . Compt. Rendus Acad. Sciences URSS, 193 2 : 42-44. 1938. Etudes sur l’histoire geologique et de phylogenie des ordres des insectes (Pterygota). Trav. Inst. Paleont., 7: 1-149. Scudder, S. H. 1893. Insect fauna of the Rhode Island coal field. U. S. Geol. Surv. Bull., 101: 1-21. Simpson, G. G. 1961. Principles of animal taxonomy. Columbia Univ. Press, pp. 1-247. AUSTRALIAN CARABID BEETLES IV. LIST OF LOCALITIES, 1956-1958* 1 By P. J. Darlington, Jr. Museum of Comparative Zoology, Cambridge, Mass. This is a brief account, with list of localities, of a collecting trip that I made to Australia from December 1956 to June 1958. My wife and fourteen year old son accompanied me. My own travel ex- penses were paid in part by a fellowship of the John Simon Guggen- heim Memorial Foundation. My purpose was to collect carabid beetles in the wetter forests of the whole eastern edge of Australia, from southern Tasmania to northern Cape York. The present paper summarizes our itinerary and then lists the localities at which Carabi- dae were obtained. I had collected at some of these localities previous- ly, in 1931-1932, as a member of the Harvard Australian Expedition. We reached Sydney December 2, 1956, and proceeded to Tasmania December 7. We bought a used (and abused) Volkswagen pickup truck in Hobart and traveled and lived in it most of the time there- after. We stayed in Tasmania until April 5, 1957. Collecting was done mostly in the wetter forests there; first into the accessible edges of the almost impenetrable southwestern forest (Arve River, Hartz Moun- tain, Mt. Field, Florentine River, Zeehan, etc., and later Corinna) ; then in mountainous north-central Tasmania (Lake St. Clair, Waldheim and Cradle Mt., Great Lake, Waratah, etc) ; and finally on isolated mountains and plateaus in the northeast (Mt. Ben Lo- mond, Blue Tier, Alt. Barrow, etc.). Although the most extensive wet forests are in the south and west, we found some of the finest stands of N othofagus (southern beech) on and near the Blue Tier in northeastern Tasmania. Lake St. Clair was the best (most diverse and richest) Tasmanian locality visited. The most southern locality reached was Cockle Creek, about 5 miles from the southernmost tip of land. On the mainland of Australia, beginning in April, 1957, we worked the Otway Ranges southwest of Melbourne, then crossed the mountains via Alt. Hotham and Omeo to Canberra, which we reached Alay 1, and where we made winter quarters in a rented house. ’Published with the aid of a grant from the Museum of Comparative Zool- ogy at Harvard College. I I I I 12 Psyche [December At Canberra I studied the types of Australian Carabidae in the Sloane Collection, now in care of the Commonwealth Scientific and Industrial Research Organization. During this time we collected repeatedly on the Brindabella Range west of Canberra, briefly on the lower slopes of Mt. Kosciusco, and at other places within reach. From Canberra, starting September i, 1957, we crossed by the Sturt Highway to Adelaide, where I examined Sloane’s and Lea’s Tasmanian carabid types. We then followed the coast south, east, and north, via Melbourne, to Sydney, making side trips to collect on Mt. Baw Baw, Wilsons Promontory, etc. From Sydney, October 15, 1957, we started a long zig-zag through northern New South Wales, going west almost to Nyngan, back to the Mt. Royal Range (inch Barrington Fops), and north via the Combovne, the Dorrigo-Ebor plateau (including New England Na- tional Park and Point Lookout, a magnificent, mile-high collecting place), Armidale, and a series of localities across extreme northern New South Wales, to Brisbane in Queensland. We reached Brisbane November 12, 1957, and left on the 1 6th for North Queensland. Going north, we worked (among other places) the Eungella Range (an important, isolated rain forest locality in- land from Mackay), and Mt. Spec, Mt. Fox, and the Kirrama Range. We reached Cairns December 6. From Cairns, we began work on the Atherton Tableland, in the largest and richest tropical rain forest area in Australia. We found the ground at Kuranda too dry, so moved our headquarters to Ather- ton and collected thereabouts, especially on the Dividing Range south and west of the town. December 13-16, 1957, we camped and collected on Mt. Bartle Frere. December 20, we returned to Ca'rns, and in the next few days did Thornton Peak (Mt. Alexandra) northeast of Daintree and Mt. Lewis southwest of Mossman. From December 31, 1957 to January 3, 1958, I collected alone on Mt. Bellenden Ker. On January 7, 1958, we flew to Thursday Island and crossed to the tip-of-peninsular forests of Cape York, collecting at Bamaga and Lockerbie until the 20th. From Lockerbie we worked to within 10 miles of the northern tip of continental Australia. My son and I interrupted the return Right at Iron Range for a week’s collecting, including a side trip on foot to Tozer Gap. We spent most of February, 1958, on the Atherton Tableland, where, however, collecting was hampered by continual heavy rain. 1960] Darlington — Carabid Beetles 1 13 From late February until March 13 we were moving south, re-visiting the K irrama Range and Mt. Spec, and reaching two additional, important localities: the isolated rain forest on the Elliot Range southeast of Townsville, and Mt. Jacob near Many Peaks, where is the northernmost rain forest (or semi-rain forest) of the South Queensland forest system. From Brisbane, March 19 to April 12, 1958, we made a loop south to Victoria, to collect on high mountains that had been snow covered before. We reached highest altitudes on Mt. Kosciusco, then drove south on the Bonang Highway into Victoria (this road cuts some fine and apparently unworked collecting areas), then collected on the summit ridge of Mt. Baw Baw, on Mt. Buller briefly, on Mt. Hotham again, and at Pretty Valley on Mt. Bogong. Returning north we touched Exeter and Ourimbah (type localities) in south- central New South Wales, Bellangry Forest northwest of Wauchope, Point Lookout again, and localities in the extreme north of New South Wales. On April 19, 1958, we started another loop, through South Queens- land, working rain forest areas, the most important being on the Bunya and Blackall Ranges. Then, beginning May 6, we spent a week at Binna Burra on the McPherson Range on the Queensland- New South Wales border. Beginning May 23, 1958, we flew back to Cairns, collected along the Black Mt. road north of Kuranda, flew halfway up the Cape York Peninsula to Coen, and from there were taken by Lea Wassell to Silver Plains and the Rocky Scrub (rain forest) on the east side of the peninsula. Returning from this trip, we stopped for nearly a week at Shiptons Flat and Mt. Finnigan, about 25 miles south of Cooktown. We were back in Sydney June 17, 1958, and from there circled through the southern Blue Mts. (Springwood) , the Burragorang Valley, Burrawang, etc. We started the flight home from Sydney June 25, reaching Boston the afternoon of June 29, 1958. All specimens collected reached Boston safely, and have been mounted and labeled and are ready for study. We got little except Carabidae and a useful series of ants collected by my son. We had neither time nor strength for general collecting. Psyche, 1960 V of,. 67, Plate 12 Map of Queensland with Inset of Atherton Tableland SOUTH AUSTRALIA Psyche, 1960 Vol. 67, Plate 13 Maps of New South Wales, Victoria, and Eastern South Australia and of Tasmania. Psyche [December 1 1 6 LIST OF LOCALITIES Localities are listed approximately from north to south, because faunal transitions are best treated in that direction, although our collecting (and the numbering of our maps) was in general in the other direction. Letters and numbers following the names refer to the accompanying maps (Q = Queensland map, etc), on which the approximate position of localities is shown by number. One number is sometimes used to include two or more adjacent localities. Names are spelled according to Gazetteer No. 40, AUSTRALIA, prepared by the United States Office of Geography, Dept, of Interior (1957), with my original spelling indicated in parentheses if different. Lati- tudes and longitudes of most localities can be obtained from this gazetteer. Topography of each locality is indicated by “low” (low- lands, inch interior plains with lowland topography even if at moderate altitudes), “plat” (plateaus rising from lowlands), “mts” (mountains), and “rge” (mountain range). Altitudes given are usually the approximate or prob(able) altitudes at which collecting was done rather than exact altitudes of fixed points. Types of forests chiefly collected in are indicated as savannah w(oodland), sclerophyll f( orest), and rain f( orest). This is the primary classification of forests in the wetter parts of eastern Australia (see “The Australian Environment,” Commonwealth Scientific and Industrial Research Organization, Melbourne, second ed., 1950, p. 78). Savannah wood- land has trees (usually eucalypts) widely spaced, and ground covered with grass or herbs. Sclerophyll forest has trees closer, and dense undergrowth of shrubs. Rain forest has trees close, often many vines, but scanty undergrowth. Of course there is variation and transition within this main classification. Savannah woodland varies in height and spacing of trees and (in Tasmania and southern Australia) some- times merges with sclerophyll forest. Sclerophyll forest may be dry or wet, and wet sclerophyll is sometimes difficult to distinguish from poor rain forest. Tropical rain forest (often with giant fig trees etc.) and south temperate rain forest (often with antarctic beeches of the genus Nothofagus) are very different botanically, and entomologically. And tropical rain forest varies considerably and (as here used) sometimes grades into seasonally dry monsoon forest. My classification of forests in the following list is somewhat arbitrary and is based on general impressions rather than exact data. Dates are those of my collecting. Previous collecting done at the same localities in 193 1- 1932 is indicated in parentheses. Single words or phrases are used 1960] Darlington — Carabid Beetles 1 1 7 to indicate principal collecting methods used at each locality. “Logs” means under logs on the ground, or in rotting logs. “Stones” means under stones. “Waterside” means beside, or in vegetation in, standing or running water. “Drowning” means throwing large quantities of loose soil, leaf litter, or debris into water and catching the Carabidae that come to the surface of the wet materials. This is a little known but very effective collecting method. “Bark” means collecting under loose slabs and flakes of bark on trunks of living trees, usually eucalyptus. A whole special insect fauna lives in this habitat in Aus- tralia. “Arboreal” means beating and sweeping. And “light” means collecting insects attracted to artificial light at night. “Usual ground m(ethods)” means first four methods listed above. “All usual m(ethods)” means all methods listed except light, which is separately specified. Lockerbie, Q41 ; low; light rain f, savannah w; Jan. 11-15, 58; all usual m, light. Bamaga, Q40 ; low; savannah w, light rain f; Jan. 7-11. 15-20, 58; all usual m, light. Iron Range, Q39; low; rain f, savannah w; Jan. 21-28, 58; all usual m. Tozer Gap, Q39; mts to over 1000 ft.; rain f, savannah w; Jan. 25-26, 58; logs, stones. Coen, named on Q map; low; savannah w, gallery f; May 27, June 4-5, 58 (also 1932) ; all usual m. Rocky River (Rocky Scrub), Q38; low to mts c. 2000 ft.; rain f., savannah w; May 30-June 2, 58 (also west side this “scrub’’ in 1932) ; all usual m inch much drowning of ground debris from upper banks of river in rain f. Silver Plains, Q37 ; low; savannah w; May 27-June 4, 58; logs, waterside, light. Cooktown, named on Q map; low; poor rain f, savannah w (much cleared) ; June 5, 10-12, 58; little collecting. Shiptons Flat, 036; c. 1000 ft.; good savannah w to gallery rain f, with transitions; June 5-10, 58; all usual m. Mr. Finnigan, Q36; 1000 to over 3000 ft.; rain f, savannah w, transitions; June 5-10, 58; all usual m (collected up to and in edge elfin f, but catch inexplicably poor). Thornton (’s) Peak (or Mt. Alexandra), Q35; c. 1000 to c. 4500 ft.; rain f, elfin f near summit; Dec. 21-22, 57; stones, logs (collecting cut short by bad weather) . Mossman-Daintree area, Q33; low; diverse vegetation (much altered); Dec. 20-27, 57; little collecting except on Thornton Peak (see above) and Mt. Lewis (see below). Mt. Lewis, Q34; to c. 4000 ft.; rain f; Dec. 25-27, 57; stones, logs. Cairns, named on Q map; low; diverse vegetation (much altered) inch remnants rain f; various dates Dec. 57-Feb. 58, May-June 58; all usual m, light. Kuranda, Q30, and named on Atherton Tab map; on plat c. 1000 ft.; rain f, some savannah w; various dates Dec. 57 to May 58; all usual m. 1 1 8 Psyche [December Atherton Tableland, Q29, and inset on Q map; plat c. 2000-3000 ft., with mts to 4000 (attached Mts. Bartle Frere, Bellenden Ker over 5000) ; many tracts rain f, good savannah w; several visits, Dec. 57 to May 58 (also 1932) , intensive collecting at various points, all usual m. Mt. Molloy (town), Atherton Tab map 12 (but town not on the tableland) ; low; dry savannah w; Dec. 27, 57; little collecting. Mareeba, named on Atherton Tab map; low; savannah w; several brief visits, but collected especially some miles north, Feb. 58; collecting especial- ly on margins shallow, flooded ponds (flood collecting). Davies Creek road, Atherton Tab 8; mts to c. 3000 ft.; good savannah w to good rain f; Dec. 27-29, 57, May 25, 58 ; logs, stones. Mountains north of Kairi, Atherton Tab 7; to 4000 ft.; rain f; Dec. 19 ,57; logs, stones. Atherton (named on Atherton Tab map) ; on plat prob. c. 3000 ft. (moun- tains south and west to 4000 ft.), good savannah w, patches rain f on flat tableland near town, much rain f on mts.; Dec. 10-20, 57, Feb. 3-11, 58; all usual m. Lake Barrine, Atherton Tab 6; on plat prob. c. 2000 ft.; rain f; no collecting this trip (much in 1932). Lake Eacham, Atherton Tab 5; on plat prob. c. 2000 ft.; rain f; Feb. 58; stones, waterside. Herberton, named on Atherton Tab map; prob. c. 2000 ft.; savannah w (prob. some rain f to east, not visited by us) ; Feb. 58; collecting chiefly in piles of dead leaves under small fallen eucalypts. Longlands Gap, Atherton Tab 3; mts prob. c. 3000 ft.; rain f; Dec. 18, 57, Feb. 11, 58; all usual m. The Crater, Atherton Tab 3; prob. over 2000 ft.; rain f; Dec. 18, 57; logs, stones. Mt. Bellenden Ker, Q32, Atherton Tab 10; to c. 5000 ft.; mostly rain f; Dec. 31, 57-Jan. 3, 58; stones, logs. Mt. Bartle Frere, Q31, Atherton Tab 9; to c. 5000 ft.; mostly rain f; Dec. 13-16, 57; stones, logs. Millaa Millaa, named on Atherton Tab map; on plat prob. c. 2500 ft.; rain f; Dec. 18, 57 (also 1932) ; stones, logs. Mt. Fisher, Atherton Tab 2; to c. 4000 ft.; rain f; Feb. 12, 58; stones, logs. Ravenshoe, named on Atherton Tab map; on plat prob. c. 3000 ft.; good savannah w, rain f south and east; Feb. 11-13, 20-21, 58; logs, stones, and special collecting by edges of rain pools west of town, washing leaf debris from under thickets etc., and treading heavy aquatic vegetation over deep water in pool of river visible from road 2 or 3 miles west of town. Crawfords Lookout, Atherton Tab 1; prob. c. 1000 ft.; rain f, savannah w; Feb. 21-22, 58; little collecting. Innisfail, Q28, and named on Atherton Tab map; low; mixed vegetation (much altered); Feb. 22, 58; no collecting in town, but some in rain f, along Millaa Millaa-Innisfail road; logs, stones. Tully Falls, Atherton Tab 4; prob. 2000-3000 ft. (above falls) ; some rain f; Feb. 58; little collecting. Tully, Q27 ; low; various vegetations; passed Dec. 57, Feb. 58; little collecting. Cardwell, Q25; low; various vegetations; passed Dec. 57, Feb. 58; little collecting. Kirrama Range, Q26, to c. 3000 ft., rain f, savannah w; Dec. 4-6, 57, Feb. 22- 23, 58 ; usual ground m. 1960] Darlington — Carabid Beetles 1 19 Ingham, Q23 ; low; mixed vegetation; passed Dec. 57, Feb. 58; little collecting. Mt. Fox plateau, Q24 ; c. 2000 ft.; savannah w, some rain f; Dec. 3, 57; poor collecting. Mt. Spec plateau (inch Paluma Range), Q22 ; to c. 3000 ft.; rain f, savannah w to west; Nov. 30-Dec. 2, 57, Feb. 23-27, 58; stones, logs. Townsville, named on Q map; low; dry savannah w (much altered by man); Nov. 29-30, 57, Feb. 27-Mar. 1, 58; little collecting near town, except some at lights in outskirts. Elliot Range, Q20; to over 3000 ft.; savannah w on slopes, rain f on high ridge; Mar. 2, 58; logs, stones in rain f on “Sharp Elliot”. Charters Towers, Q21; low; dry savannah w; Mar. 3, 58 ; a little river-bank collecting where Townsville road crosses Burdekin R. Proserpine, Q19; under 1000 ft.; mixed vegetation (much altered by man) inch palm f toward Repulse Bay; Nov. 28, 57; very poor collecting (too dry), in palm f. Mackay, named on Q map; low; mixed vegetation (altered by man) ; Nov. 21- 22, 27-28, 57, Mar. 5-6, 58; little collecting near town, but some along road toward Rockhampton. Eungella Range, Q18; to c. 4000 ft.; rain f, good savannah w to west; Nov. 22-27 , 57 ; all usual m. Byfield, Q17; low; mixed vegetation (largely altered by man) incl. rem- nants approaching rain f; Nov. 18-19, 57; logs, stones, drowning along small brook in palm f. Rockhampton, named on Q map; low; dry savannah w, swamps, dense dry sclerophyll c. 50 miles west; passed Nov. 17-20, 57, Mar. 58; little collecting near the town, mostly around swamps, but some at stated distances in various directions. Many Peaks, Q16; low; savannah w; Mar. 11, 58; no collecting at town but some along road north. Mt. Jacob, Q16; rge c. 2000 ft.; rain f (or something like it), savannah w; Mar. 11-12, 58; logs, stones. Monto, Q10; low; savannah w, tract of dense dry sclerophyll f; Apr. 23-25, 57 ; logs, stones esp. in sclerophyll f. Gayndah, Q9 ; low; savannah w, formerly rain f but latter cleared; Apr. 22- 25, 57; little collecting, mostly bark, river margins. Biggenden, Q 1 1 ; low, hills; savannah w, something like rain f to southwest; Apr. 30, 58 ; little collecting, in rain f. Goodnight Forest, Q12; low; thick dry sclerophyll; May 1, 58; poor collect- ing, but a few small carabs by water holes. Gallangowan Forest, v. Manumbar, Q13; rge c. 2000 ft.; rain f, good savannah w or open sclerophyll f; Apr. 26-27, 58; logs, stones. Jimna, Q14; moderate altitude; mixed vegetation; Apr. 27, 58; little collect- ing. Blackall Range, incl. Mapleton, Montville, Maleny, Q15; to c. 2000 ft.; rain f, good savannah w or open sclerophyll f; Apr. 27-28, May 3-5, 58; logs, stones, mostly in rain f. Kenilworth, Q15; low; mixed vegetation incl. rain f ; May 2, 58; logs, stones. Bunya Mts., Q8 ; to over 3000 ft.; rain f, good savannah w or open sclero- phyll f; Apr. 20-21, 58; stones, logs. Cooyar Creek, Q7 ; low; savannah w; Apr. 19-20, 58; logs. Benarkin, Q6; low; woods approximating rain f; Apr. 19, 58; little collect- ing. 120 Psyche [December Yarraman, Q6 ; low; savannah w, woods approximating rain f; Apr. 19, 58; logs. Brisbane, named on Q map; low; savannah w; various visits Nov. 57 to June 58 (also 1932, 1943) ; little collecting near city (specimens from nearest rain f, on Mt. Glorious & Mt. Tamborine, labelled from these places) . Mt. Glorious, Q5; prob. c. 1000 ft.; rain f, good savannah w; mid-May 58; little collecting (more in Nov. 43), stones, logs, mostly in rain f. Mt. Tamborine, Q4; plat c. 2000 ft.; rain f ; not visited 1957-58 but collected Nov. 1943 ; logs, stones, drowning debris in rain f. Cunninghams Gap, Q2 ; mts c. 2000-4000 ft.; rain f; May 13-14, 58; logs, stones (most specimens taken in gap proper, but we collected from there nearly to summit Mt. Cordeaux, c. 4000 ft.). McPherson Range (Q. Nat. Park), Ql; to c. 4000 ft.; mountain rain f, some savannah w, small areas almost pure Nothofagus f on highest points; May 58 (see Binna Burra, below) (collected on McPherson Range also 1932). Binna Burra (Binnaburra) , Ql ; on rge c. 3000 ft. and over; mountain rain f, some savannah w; May 6-10, 58; usual ground m. Beechmont, Ql; on rge c. 3000 ft.; rain f., savannah w; passed repeatedly 1957-1958 but collected only 1943, in rain f probably now cleared. Springbrook, Ql ; on rge c. 2000 ft.; mixed vegetations inch rain f; May 10-11, 58; little collecting, logs, stones. Stanthrope, Q3 ; to prob. c. 5000 ft.; savannah w etc. (no rain f) ; passed repeatedly 1958; no personal collecting, but obtained specimens from Mr. E. Sutton. Wyberba, Q3 ; to over 5000 ft.; savannah w and/or good sclerophyll f, rocky and bushy country; May 16, 58; poor collecting (dry year) but good locality, specimens obtained from Mr. Sutton. Woodenbong, NSW56; moderate alt.; good savannah w; May 17, 58, and other dates; poor collecting (dry year), logs. Unumgar Forest (Cox’s Road), NSW61; mts c. 2000 ft.; mostly rain f; May 17-18, 58; logs, stones. Wiangaree, NSW57; low; savannah w; Nov. 9, 57; waterside. Murwillumbah, NSW60; low to c. 1000 ft.; mixed vegetation inch rain f c. 10 miles westsouthwest (at Byrrill Ck.) ; Nov. 10-11, 57; stones, logs, in rain f. Dunoon, NSW59; moderate altitude; mixed vegetation including rain f at rather low altitude (also rain f at prob. c. 2000 ft. up forestry road to Whian Whian Forest) ; Nov. 10, 57, Apr. 12, 58 ; logs, stones in rain f. Lismore, NSW58; low; mixed vegetation inch small piece rain f preserved in town, larger piece c. 4 miles northeast; several dates especially Apr. 11, 58 ; logs, stones, mostly in rain forest 4 miles northeast. Acacia Plateau, NSW54; c. 2000 ft.; rain f; Nov. 7-8, 57; logs, stones. Acacia Creek, NSW53; low; mixed vegetation, now no real rain f; Nov. 7, 57 ; little collecting. Tooloom Range, NSW55; to c. 2000 ft.; rain f south slope, also savannah w; Nov. 8-9, 57, May 16-17, 58; usual ground m inch drowning debris from beside small temporary pond in opening in rain f. Wilsons Downfall, NSW52; moderate altitude; savannah w; Nov. 7, 57; little collecting. Tenterfiei.d, NSW51; moderate to rather high altitude; savannah w; Nov. 7, 57 ; collecting at stated distances in various directions, logs, stones. 1960] Darlington — (Jarabid Beetles 121 Glen Innes, NSW49; various altitudes; savannah w ; Nov. 5, 7, 57, and other times briefly; collecting at stated distances in various directions, logs, stones. c. 40 miles eastnortheast (ENE) Glen Innes, NSW50; moderate altitude; savannah w, heathy f, patches rain f; Nov. 6, 57; usual ground m. Armidale, NSW47; moderate altitude; savannah w; Nov. 2-5, 57, and other brief visits; little collecting. Uralla, NSW48; moderate altitude; savannah w; Nov. 3, 57; logs. Ebor, NSW45; to c. 5000 ft.; savannah w; Oct. 31-Nov. 1, 57, Apr. 9, 58; logs, stones. New England National Park (Point Lookout), NSW46; to over 5000 ft.; rain f inch N othofagus , some savannah w; Nov. 1-2, 57, Apr. 9, 58; usual ground m. Dorrigo, NSW44; plat c. 2000-3000 ft. (and same plateau west rises to c. 5000 ft. at Ebor etc.) ; rain f, savannah w; Oct. 31, 57 (also 1932) ; stones, logs. Bellingen, NSW43; low; mixed vegetation inch remnants of something like rain f; Oct. 30-31, 57; logs, stones. The Big Hill (Kempsey-Ebor road), NSW42; up to Ebor plat; semi-rain f; Apr. 8, 58 ; logs, stones. Bellangry Forest (northwest of Wauchope), NSW41; mts to prob. c. 3000 ft.; rain f, said to include N othoj agns ; Apr. 6-7, 58; logs, stones. The Comboyne, NSW39; plat to c. 2800 ft.; mixed f inch remnants rain f in gullies; Oct. 28-29, 57; logs, stones, especially near Mt. Gibralter and in gully northeast. Port Macquarie, NSW40; low; mixed f inch small tract something like rain f; Oct. 29-30, 57; logs. Mt. Royal Range, NSW33; plat to c. 5000 ft.; savannah w on slopes, savan- nah w and rain f inch N othofagus on plat; Oct. 23-27, 57; usual ground m (see 3 following localities). Barrington House, NSW33; low at southeast foot Mt. Royal Range; gallery rain f along river; Oct. 25-26, 57; usual ground m. Barrington Tops, NSW33; plat on southeast end Mt. Royal Range, which see (above) ; Oct. 27, 57 (also 1932) ; usual ground m. Tomalla Tops, NSW33; plat on northwest end Mt. Royal Range, which see (above) ; Oct. 23-25, 57; usual ground m. Gloucester, NSW37 ; low; savannah w; Oct. 28, 57; logs. Paterson, NSW35; low; savannah w; Oct. 25, 57 ; logs. Stratford, NSW36; low; savannah w; Oct. 28, 57; logs. Booral (10 miles east of), NSW38; low; savannah w, thicker in stream gully; Apr. 6, 58; logs. Ourimbah, NSW34; lowr; savannah w, formerly semi-rain f (we did not And latter) ; Apr. 5, 58; poor collecting. Tamworth, NSYV32; low; savannah w; Oct. 22, 57; chiefly waterside. Mt. Kaputar, NSW31; to c. 4000 ft.; savannah w with underbrush in places; Oct. 20, 57; logs, stones, chiefly near summit. Narrabri, NSW30; low; savannah w and/or rather dry sclerophyll f; Oct. 19, 57 ; little collecting. Coonabarabran (north of ) , NSYV29; low; savannah w and/or dry sclero- phyll f; Oct. 19, 57; logs, stones. Nyngan, NSW27; low; originally dry savannah w or sclerophyll f, now mostly cleared for sheep; in vicinity Oct. 17-19, 57; poor collecting (dry year) at stated distances south and east. 122 Psyche [December Bogan River, NSW28; low; originally dry savannah w or sclerophyll f, now mostly cleared for sheep; Oct. 18-19, 57; poor collecting (dry year) except by edges pools in bed of Bogan R., but specimens obtained from Mr. J. W. T. Armstrong. Narromine, NSW26; low; savannah w and/or rather dry sclerophyll f; Oct. 17, 57 ; little collecting. Dubbo, NSW25; low; savannah w or rather dry sclerophyll f; Oct. 17, 57; little collecting. Orange, NSW24; low; originally savannah w or rather dry sclerophyll f; Oct. 16, 57; little collecting. Mt. Canobolas, NSW24; to c. 4500 ft.; savannah w; Oct. 16-17, 57; stones, logs near summit and especially near camp site at base (specimens taken at base are so labeled). Blue Mts., NSW21; mts to over 3000 ft.; various vegetations inch bushy heath, sclerophyll f, savannah w, wetter f in ravines; collected only at Springwood (see below) in 1957-58 (but at several localities in 1932). Springwood, NSW22, prob. c. 1000 ft.; savannah w, wetter f in ravine to south; June 58; logs, stones, especially in wet ravine. Burragorang Valley, NSW23; low; savannah w; June 58; logs. Sydney, named on NSW map; low; originally savannah w, dry sclerophyll, etc., but now much altered by man; in vicinity various dates 1956-1958 (also 1931-1932, 1943) ; little collecting near city. National Park, NSW20; low; sclerophyll f etc.; Oct. 9-10, 57 (also 1931) ; little collecting. Exeter, NSW15; low; savannah w; Apr. 4, 58; logs. Burrawang, NSW19; low; savannah w; June 58; logs, stones. Macquarie Pass, NSW18; edge plat; savannah w and thickets; June 58; drowning ground debris. Jamberoo, NSW17; low; savannah w and wetter thicker f in ravine; June 58 ; stones, logs in wetter f. Kiama, NSW16; low but Saddleback Mt. to prob. c. 2000 ft.; savannah w much cleared, but remnants of good woods still on Saddleback; Oct. 8-9, 57; logs, stones in f on Saddleback. Saddleback Mt., see Kiama, above. Termeil (Batemans Bay), NSW13; low, but worked into hills northwest nearly to Pigeon House; savannah w; Oct. 5-7, 57; logs, stones, and by pools intermittent brook in hills toward Pigeon House. Batemans Bay, see Termeil (above). Pigeon House, see Termeil (above). Clyde Mt., NSW14; prob. c. 2000 ft. (edge plat) ; good mixed forest with considerable underbrush ; several visits from Canberra May-Aug. 57 ; logs, stones. Braidwood, NSW12; low (but on plat); savannah w (mostly cleared); July 11, 57 ; flood collecting along river and streams. Moruya, NSW10; low; savannah w (mostly cleared); July 11, 57; flood collecting. Bodalla, NSW9; low; savannah w (mostly cleared) ; July 11, 57; collecting esp. by standing flood water in woods. Brown Mt., NSW11; prob. c. 2000 ft. (edge plat); good woods with con- siderable underbrush ; Oct. 4, 57 ; logs, stones. Bega, NSW8; low; savannah w (partly cleared) ; Oct. 4, 57; logs, stones. Eden, NSW7; low; savannah w (much cleared); Oct. 4, 57; collecting at stated distances north and west, logs, stones. Cooma, NSW3; on plat; savannah w; passed several times; little collecting. 1960] Darlington — Carabid Beetles 123 Adaminary Dam, NSW2; on plat; savannah w; Aug. 3, 57; attempted flood collecting as dam filled, but unsuccessful. Mt. Kosciusco, NSW1; to over 7000 ft. (highest mt in Australia); good sclerophyll f on slopes, open snow gum w higher, mountain moors still higher; Aug. 3, 57, Mar. 22-23, 58 (also 1931) ; usual ground m, bark. Albury, NSW4; low; savannah w etc. (mostly cleared) ; passed on various occasions; little collecting. Balranald, NSW6; low; various dry w; Sept 2-3, 57; little collecting. Narrandera, NSW5; low; various more or less dry w; Sept. 1-2, 57; little collecting, mostly by drowning broken earth Canberra, Australian Capital Territory, named on NSW map; in wide valley on plat; savannah w; May through Aug., 57; usual ground m, bark, but collecting poor (dry year). Brindabella Range (inch Mt. Franklin and Mt. Gingera), ACTl on NSW map; to over 6000 ft.; savannah w, sclerophyll f, open snow gum forest (in order with increasing altitude) ; May, June, 57; usual ground m. Gudgenby River, ACT2 on NSW map; in valley on plat; savannah w; June 9, 57; logs, stones, bark. Mixta Mixta, V30; low; sclerophyll f; Apr. 30-May 1, 57; little collecting. Glen Wills, V29; moderate altitude; sclerophyll f; Apr. 30, 57; bark. Omeo, V28 ; low; sclerophyll f; Apr. 25-30, 57; logs, stones, bark, especially along road toward Mt. Hotham. Mt. Bogong inch Pretty Valley, V27; plat to c. 6000 ft.; sclerophyll f on slopes, open snow gum w and moors higher; Apr. 2, 58; logs, stones. Pretty Valley, see Mt. Bogong. Mt. Hotham, V26; to c. 6000 ft.; sclerophyll f on slopes, open snow gum w and moors higher; Apr. 24-28, 57, Mar. 29-31, 58; usual ground m, bark. Bright, V 3 1 ; low; sclerophyll f; Apr. 23-24, 57, Mar. 29, Apr. 1, 3 (toward Mt. Buffalo on latter day), 58; logs, stones, bark. Mt. Buli.er, V25; to over 5000 ft.; savannah av, sclerophyll f on lower slopes, open snow gum av higher; Mar. 28, 58; logs, stones, drowning debris from under clumps of snoAV gums. Jamieson, V24; moderate altitude; sclerophyll f; Mar. 27, 58; little collecting. Woods Point, V23; moderate altitude; sclerophyll f; Mar. 27, 58; little collecting. Aberfeldy, V22; moderate altitude; savannah w and/or sclerophyll f; Mar. 27, 58 ; logs, stones. Walhalla, V21 ; moderate altitude; sclerophyll f and/or savannah w; Mar. 27, 58 ; little collecting. Mt. Baw Baw, V9; to c. 5000 ft.; sclerophyll f on slopes, open snow gum w higher, vestiges rain f inch Nothofagus in gullies c. 4000 ft.; Sept. 27-28, 57, Mar. 25-27, 58; logs, stones on wooded slopes (1957), and in high snoAV gum w (1958), and drowning debris from Nothofagus gully (1958) (all “Trechus” by last method). Mt. Donna Buang, VI; to c. 4000 ft.; sclerophyll f, approaching temperate rain f in gullies; several visits Apr. 57 (also 1931) ; logs, stones, drowning, especially near summit and at Cement Creek. Cement Creek, see Mt. Donna Buang. Bonang Highway, V20; moderate altitudes (from plat south to low); sclerophyll f approaching temperate rain f in gullies; Mar. 24, 58; logs, stones, droAvning. Mt. Drummer, V19; prob. to c. 2000 ft.; savannah w, sclerophyll f; Oct. 3, 58 ; logs, stones, especially in gully north of road on east slope. 124 Psyche [December Orbost, V18; low; savannah w and/or sclerophyll f; Oct. 2-3, 57; little collecting. Lakes Entrance, V17 ; low; savannah w and/or sclerophyll f; Oct. 2, 57; little collecting. Bairnsdale, V16; low; savannah w and/or sclerophyll f; Oct. 2, 57; little collecting. Stratford, V 1 5 ; low; savannah w and/or sclerophyll f; Oct. 2, 57; little collecting. Sale, V14; low; savannah w; Oct. 1-2, 57; flood collecting. Yarram, Vl 3 ; low; savannah w and/or sclerophyll f; Oct. 1, 57; little collecting. Tarra Valley, V12, on rge prob. c. 1000 ft.; sclerophyll f approaching tem- perate rain f in wet gullies; Sep. 30-Oct. 1, 57; logs, stones. Gunyah, Vll; on rge prob. c. 1000 ft.; sclerophyll f approaching temperate rain f in wet gullies; Sept. 30, 58; logs, stones. Wilsons Promontory (most southern point of continental Australia), V10; low; sclerophyll f, heath, moors, dunes; Sept. 28-30, 57; poor collecting, tried especially in forest at Lilly Pilly Gully. Noojee, V8 ; moderate altitude; savannah wr to sclerophyll f; Sept. 26, 57, Mar. 25, 58 ; logs in woods by Neerim-Noojee road. Melbourne, named on V map; low; savannah w (mostly cleared) ; Apr. & Sept., 57 ; little collecting near city. Werribee Gorge, near Bacchus Marsh, V7; low; savannah w; Sept. 19, 21, 57; logs, stones, waterside (beautiful locality, but dry year). Beech Forest, see Otway Ranges. Otway Ranges inch Beech Forest, V2; plat c. 2000 ft. ; sclerophyll f ap- proaching rain f in ravines, some N othofagus ; Apr. 12-13, Sept. 17-18, 57; logs, stones drowning, especially at Tanybryn Junction and Crowes. Port Fairy, V6; low; various vegetations; Sept. 17, 57; little collecting. Portland, V5; low; various vegetations; Sept. 17, 57; little collecting. The Grampians, V3; mts prob. to over 2000 ft., dry sclerophyll w, heath; Apr. 14-15, 57; stones, logs (poor collecting, dry year). Mildura, V4; low; dry woods etc.; Sept. 3, 57; little collecting. Renmark, SAl ; low; dry woodland etc., much cleared; Sept. 4, 57; little collecting. Adelaide, named on SA map; low; savannah w (much cleared) ; Sept. 6-14, 57 (also 1931) ; little collecting near city. Mt. Lofty Range, SA2 ; plat to c. 2000 ft.; savannah w etc.; Sept. 9, 14, 57; stones, logs. Murray Bridge, SA3; low; various vegetations; Sept. 15, 57; collecting most- ly along road south toward Meningie. Meningie, SA4; low; various dry vegetations; Sept. 15, 57; collecting mostly along roads north and east. The Coorong, SA5 ; lagoon connected with ocean ; various dry vegetations, sand; Sept. 15, 57; little collecting. Kingston, SA6; low; various dry vegetations; Sept. 16, 57; little collecting. Princes Highway at South Australia-Victoria line; V7 ; low; good savan- nah w; Sept. 16, 57; logs, digging (carenums and Teropha). Launceston, named on T map; low; savannah w etc.; based here Feb. 24- Apr. 5, 57 ; little collecting near city. Mt. Barrow. T26; to c. 4000 ft.; sclerophyll f on slope, N othofagus thickets near tree line, open summit; Apr. 2, 57; stones near summit, drowning debris from N othofagus thickets, logs, stones in f near base. i960] Darlington — Carabid Beetles 125 Mt. Ben Lomond, T27 ; to c. 5000 ft.; savannah w near base, sclerophyll f on slopes (no rain f but ti tree flats and wet gullies in sclerophyll area), moors etc. above c. 4000 ft.; Mar. 5-10, 57; usual ground m, bark. Blue Tier (plat), T28 ; plat c. 2000 ft.; temperate rain f inch Nothofagus (partly cleared) ; Mar. 26-Apr. 1, 57; usual ground m inch much drowning of debris from rain f. Included in collection from this locality are speci- mens taken in Nothofagus f (big trees) beside road between Goshen and Weldborough. Lottah, T28 ; probably c. 1000 ft. (foot of Blue Tier); probably formerly sclerophyll f; Mar. 26, Apr. 1, 57, little collecting. This is an old type locality, but the vegetation has been much altered, and the old collections may have been made many miles from the town of Lottah, perhaps on the Blue Tier. Goulds Country, T28 ; moderate altitude; wet sclerophyll f; Apr. 1, 57; logs, stones, drowning debris from under thickets. Mole Creek, T25; low; savannah w; Apr. 3, 57; bark, and unsuccessful attempted cave collecting. Mersey River, T19; deep river valley; sclerophyll f; Mar. 2-3, 57; poor collecting, mostly washing river banks. Waldheim, v. Cradle Mt., T17; mts c. 4000 ft.; temperate rain f inch Nothofagus , open snow gum w, moors; Feb. 23-Mar. 1, 57; usual ground m inch much drowning of debris from rain f and from under clumps of snow gums. Cradle Mt., T18; c. 5000 ft.; moors, bushy heath, rocks, mountain lakes; Feb. 26, 57 ; stones to c. 4500 ft., a little drowning. Burnie, T20 ; low; savannah w; Mar. 13, 23, 57; little collecting. High land south of Burnie, T21; prob. c. 1000-2000 ft., wet sclerophyll f, temperate rain f inch Nothofagus ; Mar. 20-23, 57; usual ground m. Waratah, T22 ; prob. c. 2000 ft.; temperate rain f inch Nothofagus , wet sclerophyll f; Mar. 13-14, 18-20, 57; usual ground m inch much drowning of debris from rain f. This is a famous old type locality. For practical purposes it includes Magnet (not visited by us), which is nearby. Magnet, T22, see Waratah, above. Whyte River, T23 ; low; sclerophyll f, temperate rain f inch Nothofagus ; Mar. 17-18, 57 ; usual ground m. Corinna, T24; low; temperate rain f inch Nothofagus ; Mar. 14-17, 57; usual ground m. Zeehan, T14; low; wet sclerophyll f, rain f in places (but much cleared) ; Jan. 17-19, 57; usual ground m, mostly c. 8 miles north, and along Rosebery Road. Queenstown, T13; low; vegetation stripped; Jan. 17, 19, 57; little collecting except along road toward Lake St. Clair. (The) King River, T12; low; original f stripped, now ti tree thickets etc.; Jan. 17, 19, 57; mostly washing river bars. Lake St. Clair, Til; plat and mts c. 2400 ft. and over; savannah w (to south), wet sclerophyll f, heavy temperate rain f inch much Nothofagus ; Jan. 11-17, 19, 57; usual ground m, and waterside near lake, treading wet moor toward Mt. Rufus. Great Lake, T15; on plat c. 3500 ft.; savannah or snow gum w, open (stripped?) country, moors; Jan. 20 (south end of lake), Mar. 4 (north end), 57; stones, logs at south end, drowning debris from under pencil pines above north end of lake. Ouse Valley, T16; deep valley; savannah w; Jan. 20, 57; logs etc., bark. 126 Psyche [December Tarraleah, c. 20 miles southeast L. St. Clair (Til); moderate altitude; sclerophyll f (also rain f?) ; Jan. 11, 57; little collecting (but specimens received from Prof. V. V. Hickman). Ellendale, T10; low; savannah w and/or sclerophyll f; Jan. 11, 57; water- side (by pond in open). Mt. Field, T8 ; to c. 4000 ft.; wet sclerophyll f, temperate rain f on slopes, open snow gum w, moors above, mountain lakes; Jan. 2-7, 57; usual ground m especially logs, stones toward Mt. Field West, drowning debris from under pencil pines near Lake Dobson, drowning in temperate rain f c. half way up road, logs, bark at base. Florentine Valley, T9; low; wet sclerophyll f with gigantic eucalypts, strips of rain f with Nothofagus ; Jan. 25-27, 57; usual ground m. Hobart, named on T map; low; savannah w etc.; based here Dec. 7, 56 to Feb. 23, 57 ; specimens labeled “v. Hobart” are from ^ ,. „ rv f/& A V. TFT ^ "V/ ^ % cp ^ F"’ ^^yS..^»- % t fefejlltl f v feWr "VflaJj <- mw •/’ I :#v \ ^ ?“ — . V j?A>> .-*• -Sta.jkVi '- ?#?&X FMf| Pfeff XC X o ,J?' •%