Bert Hölldobler.
Communication, Raiding Behavior, and Prey Storage in Cerapachys (Hymenoptera: Formicidae).
Psyche 89(1-2):3-23, 1982.

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PSYCHE
Vol. 89 1982 No. 1-2
COMMUNICATION, RAIDING BEHAVIOR
AND PREY STORAGE IN CERA PACHYS
(HYMENOPTERA; FORM ICIDAE)*
BY BERT HOLLDOBLER
Department of Organismic and Evolutionary Biology, MCZ - Laboratories
Harvard University, Cambridge, Mass. 02138 U.S.A. The former subfamily Cerapachyinae was recently recognized by Brown (1975) as a tribe (Cerapachyini) within the subfamily Poneri- nae. All of the cerapachyine ant species investigated feed entirely on ants (see review in Wilson 1958: Brown 1975). During foraging cerapachyine workers engage in mass expeditions during which they raid the nests of the prey species, capturing preferably larvae and pupae, but also occasionally adults and returning them to the raid- ers nest.
Although the detailed field observations on cerapachyine forag- ing raids reported by Wilson (1958) strongly suggest that the raiding expeditions follow chemical trails, this has not yet been experimen- tally investigated. In fact, almost nothing was hitherto known about the behavioral organization of the raiding expeditions and the under- lying communication mechanism. This paper presents the first ex- perimental analysis of the raiding behavior of a cerapachyine ant species. MATERIALS AND METHODS
Three colonies of Cerapahs (?) turneri (turneri group) (acces- sion #163a, b, c; voucher specimens in Australian National Insect *Manuscript received
the editor January 22, 1982.
3




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4 Psyche [VOI- 89
Collection, ANIC, Canberra) were collected from nests in the so i 1 in a sclerophyl scrub pasture near Eungella, North Queensland (A us- tralia). One colony had a single ergatoid queen; the other co lo nies had two ergatoid queens apiece. Each colony was housed in sepa rate glass tube nests (8cm X 0.6cm ^>), with water trapped at the bo t t o m s behind cotton plugs. Each nest tube
was placed into arenas of
varying sizes, depending on the experimental design. Histolo~ical studies were conducted according to the procedures described in Holldobler and Engel 1978. Additional methodological details will be given with the description of the individual experiment, as pre- sented below.
Raiding behavior and paral-ysis of prey larvae Species of the genus Cerapachys seem to preferably prey on ant
species of the myrmicine genus Pheidole (Wilson 1958; B r o w n 1975). When 1 provided Cerapachys with colonies or fragment s of
colonies of a variety of species of the genera Iridom.~-rmex, Meran op- lus, Monomorium, Crematogaster, Pheidole, Stigtnacros, Pol\^t-ha- chis, Camponotus (placed in a 65 X 120 cm arena) they preyed freely only on Pheidole. They also accepted Monomorium larvae as p x-ey, but only when these insects were directly inserted into the e r a - pa* nest. When the Cerapachys workers encountered worker- s of the other species, or came close to their nest tubes, they usu ally showed avoidance behavior. The reaction was very different, how- ever, when individual scouts of Cerapa*~ discovered the nest t ube of Pheidole (accession #209, voucher specimens in ANIC). The C e r a - pachys worker vigorously vibrated its short antennae and m o v e d slowly into the nest tube, which contained approximately 200 PAei- dole workers and soldiers and about 150 larvae and pupae. It did
not venture very far into the foreign nest but left after a short while and ran, in a somewhat meandering route, back to its own nest, located 70cm away from the Pheidole nest. During homing it appeared frequently to touch the ground with its abdominal tip, as if it were laying a chemical trail or depositing scent spots. Seconds after it had entered the nest of its own colony, its ndstmates beca me very excited. Many grouped around the scout ant, which repeatedly raised its gaster upwards. Within one minute the scout left the nest



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19821 Holldobler - Cerapachys 5
again and moved in direction toward the Pheidole nest tube. It was closely followed by 17 nestrnates. The leading scout ant continued to move with its abdominal tip close to the ground, but intermittently it paused or moved much slower while raising its gaster slightly upwards (Fig. 1). When the Cerapachys column arrived at the Phei- dole nest tube they invaded it and attacked the Pheidole workers and soldiers. Pheidole fought back but without any effect. The heav- ily scierotized and specially protected Cerapachys (Fig. 2) were not at all affected by the mandibular grip of the Pheidole soldiers, even when they were attacked simultaneously by 3-5 Pheidole (Fig. 3). Although Pheidole outnumbered the Cerapachys invaders more than 10 times, they were rapidly disabled by the obviously very Figure 1, Recruiting C~r~pa&r~ worker. (a) Worker walking with its abdomi- nal tip ctmc 10 the ground.
(b) Worker raising the gasier upwards; arrow indicates the poiiit~on of the opening of the pygidial giand.



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6 Psyche [Vol. 8g
Figure 2.
Longitudinal section through the head and part of ihe thorax (a) and through part of the petiolus and gaster (b) of a Cerapachvs worker. Arrows indicate cuticle projections over intersegmental membranes (IM),



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effective stinging attack of the Cerapa-, during which the raiders grasped the: Pheidofe with their short mandibles, simultaneously bending their gasters forward, so that in each case the tip. where the sting extrudes, touched the opponent's body. Each sequence usually lasted less than 1 second. Almost immediately after such an attack the Pheidofe appeared to be immobili7ed. Only a few Pheiciok workers escaped from the nest tube into the arena, some of them carrying brood. After approximately I5 minutes almost all Phwioie adults in the nest tube were disabled or kilied but not a single Cerapachys worker was dead or visibly injured. Next the Cera- pachys began transporting the dead and immobilized Pheiiiote adults to their own nest. After the first workers of the raiding expe- dition had returned and unloaded the booty they returned to the Pheidole nest. Some of them raised the gaster repeatedly upwards, upon which several additional Cerapacbys workers followed them to the Pheidole nest, where they participated in the retrieval of the prey. Only after most of the Pheidole adults had been retrieved did the Csrapachys begin to transport the Pheidole brood. Each larva and pupa was briefly stung before it was picked up and carried to the Cerapathys colony. Interestingly, after approximately half the brood had been retrieved, Cerapachys nest workers began discard- ing all the dead and disabled Pheidole adults, and the next day only



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8 Psyche [VOI. 89
Pheidole brood was stored in the Cerapachvs nest. Apparently the booty of this raiding expedition was so abundant that Cerapachys preferred to keep only the more valuable and better preservable brood of the prey species, and they discarded the less valuable cadavers of the adult Pheidole. In other instances, however, where Cerapams had only adults of prey species available, I observed Cerapam feeding on the gasters of dead Pheidole workers and soldiers.
This experiment was conducted on the 25th and 26th of October 1980. At this time there was no Cerapachvs brood in the colony. On November 10, 1980, 1 noticed the first large clutch of eggs in the Cerapachys nest tube. On December 11, 1980, the colony had many large (presumably last instar) larvae, and another large cluster of eggs (Fig. 4). The colony still contained a very good supply of Pheidole larvae (Fig. 4), which did not grow or develop further but which were obviously alive. Under the microscope one could see that the prey larvae slightly moved their mouthparts. Workers, queens and larvae of Cerapa*~ all fed on the Pheidole larvae. On December 26, 1980, there were still some prey larvae left. Many of the large Cerapachys larvae had pupated; in addition the nest con- tained many medium sized larvae and another large clutch of eggs. On January 3, I98 1, a Cerapachvs worker was observed leaving the nest tube and venturing out into the arena, for the first time since October 27, 1981. At this time I provided another fragment of a Pheidole colony with larval brood in the arena; and on January 5, 198 1, Cerapachys conducted another raid, very similar in details to that just described. The fact that the captured Pheidole larvae were kept alive inside the Cerapachys nest chamber for a period of more than two months (but did not pupate or visibly increase in size) strongly suggested that they were sustained in a state of metabolic stasis. Recently Maschwitz et a1 (1979) provided experimental evi- dence that the ponerine species Harpegnathus saltator and Lepto- gen.13.\ chinensis paralize prey objects by stinging and thereby are able to store prey a limited time. In one case the preserving paralysis effect was observed to last for two weeks, and in no instance did the stung prey object ever recover from the paralysis. Similar observa- tions have been made independently by Traniello (unpublished data) with the ponerine species A mblyopone pallipes.



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Figure 4.
Fractions of a Cwaparhvs colony, with paralysed prey larvae. Q: erga-
toid queens; E: eggs; C: Cerapwhvs larvae; P: Phercfde prey larvae.



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10 Psyche [VOI. 89
As just noted, CerapaMs workers apparently sting each Phei- dole larva and pupa during the raid, before they transport the vic- tims to their nest. This appears to be a very stereotyped behavior. For example when I shook a Cerapach.rs colony which contained Pheidole larvae out of the nest tube into the arena, so that they had to move back into the nest, Cerapachys workers picking up a Phei- dole larva almost invariably went through the typical stinging motion pattern. They did not do this, however, when they picked up their own larvae. Although stinging behavior did not frequently occur inside the nest, occasionally 1 observed a Cerapachys stinging several larvae while reshuffling a pile. The Pheidole larvae are small and tender and the powerful Cera- pachys sting (Fig. 5) could easily pierce the larva and thereby kill it. Thus the injections of a paralyzing secretion through the sting has to be very subtle in order not to kill, but to preserve the larva. Brown (1975) describes the differentiated pygidium (Fig. 6) with its denticu- late margins, being present in all workers and queens of cera- pachyine ants. Brown states that "the function of the denticle- bordered pygidial plate is not known from direct observations, but it is assumed to have something to do with helping the insects to force their way through passages and cracks in soil or rotten wood, perhaps in connection with their entry into nests of termites or ant prey species".
Our morphological and histological investigations have revealed that these denticuliform and spinuliform setae on the pygidium of Cerapachys turneri and Sphinctom.\~rmex steinheili are sensory setae and comprise probably mechanoreceptors (Fig. 7). It is most likely that during the stinging process these mechanoreceptors sig- nal the gaster tip's touch of the prey larva and the extent of the stings' protrusion is thereby regulated. Many of the nonsocial acu- leate Hymenoptera, which paralyze prey by stinging, are equipped with mechanoreceptors on the tip of the sting sheath (Oeser I96 1, Rathmayer 1962, 1978). We did not detect similar structures on the tip of the sting sheaths of Cerapachys or Sphinctomyrmex. In addi- tional experiments I further confirmed the suggestion that the prey larvae, captured by Cerapachys, are preserved alive. Approximately 30 Pheidole larvae collected from a Pheidole colony were put with- out workers in a small test tube, which was kept moist by a wet cotton plug. A second similar test tube contained 30 Pheidole larvae which were taken from the Cerapachys nest. In two replications the



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picture shows the partly extruded å´iting surrounded by ihe sensory f-etue at the ..
pygidium, and last exposed sterniie. (b) Cloiic-up or ihe two kind'i ol'setae at the pygidium.




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Psyche [Vol. 89
Figure 6.
SEM picture of frontal view of pygidium of a Cerapoc'hvfi worker (a), and a worker of Sphincitimyrnw.~ vietnheili (b). Mole the arrangement of the two kinds of setae on the iruncaied pygidial piate of both species.



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14 Psyche [VOI. 89
larvae taken directly from the Pheidole colony were all dead after two weeks. On the other hand all of the larvae from the Cerapachjls colony were obviously still alive after two weeks, many of them moving their mouthparts slightly. These findings clearly demon- strate that Cerapachjs can store living prey larvae for a considerable period of. time. This food storage system appears to enable Cera- pachj!s to stay inside their nest for longer intervals. They evidently do not conduct raids as long as a good food supply is present. The following experiments were designed to test this hypothesis. One day after the Cerapachys colony B had conducted a raid on Pheidole all prey larvae were removed. As a control I manipulated colony A in the same way, but the prey larvae were immediately returned to colony A. A few days later I observed scouts of colony B in the arena, where I had provided a nest tube with a fraction of a Pheidole colony, and within a period of 4 (test I) and 7 days (test 2) colony B had conducted another raid. In the control colony A I noticed a worker briefly leaving the nest tube only once and then without venturing far into the arena. Although a tube containing Pheidole workers and brood was also provided in the arena of colony A, this colony did not conduct another raid until its supply of prey had declined considerably.
Emigration behavior
Although it is still an open question whether the Cerapachyini are nomadic, Wilson (1958, 1971) and Brown (1975) suggested that nomadism in the ant-preying cerapachyine species could well be adaptive to avoid depleting the food supply in a given neighbor- hood, just as it is in the army ants. This assumption of a nomadic life style is further supported by Brown's observations that the nests of many cerapachyine species appear to be impermanent, and that the "brood show a strong tendency to be synchronized, like those of army ants and nomadic Ponerinae". Brown (1975) also pointed out that the larvae of the Cerapachyini have a slender and cylindrical shape (G. C. Wheeler and J. Wheeler 1964), which makes them easy to transport longitudinally under the bodies of workers in the manner of other predatory and nomadic ants, such as Eciton, Aenic- tus, Dorylus, Leptogenp and Onychomyrmex. Although I was unable to demonstrate periodic nomadic behavior of Cerapachys in



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the laboratory, I could easily initiate nest emigrations by removing the waterplug and thereby causing the nest tube to quickly dry out. Individual workers soon ventured into the arena and eventually discovered a new moist nest tube located approximately 20-3Ocm away from the 01d nest. After exploring the new nest site the scout moved back to the colony, When entering the nest tube it exhibited the same behavior as when recruiting to a raid, including a repetitive Iifting of the ester. When the scout left the nest again to return to the newly discovered nest site, it was usuaily fo1bwed by several ants. Most of these first recruits a h showed the ester raising behavior on their return to the colony, and soon the whole colony began to leave the old nest tube and move ta the new one. The lamae and pupae were carried in the manner Brown (1975) pre- dicted, dung lmgitudinaily under the bodies oithe workers (Fig. 8). Adult transport was never observed; the ergatoid queens and even relatively freshly eclosed workers moved on their own to the nest site. The colonies did not contain males. After the workers had moved most of their own brood, they transported the prey larvae



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16 Psyche [VOL 89
From the ants' orientation behavior it appeared that they were following chemical trails during the nest emigration. In fact, the recruitment behavior during nest emigrations and raiding appeared to be identical. The following experiments were designed to analyze further the communication mechanisms involved in both events. Comtnunication during emigration and raiding Two distinct behavioral patterns were observed in Cerapachj?.~ ants during recruitment. (I) They seem to lay a chemical trail when returning from the target area (prey colony or new nest site) by frequently touching the abdominal tip to the ground; and (2) when close to or just entering the nest, they repeatedly raised their gaster upwards into a "calling position" and continued to do so when they moved back to the target area, usually being closely followed by a group of recruited nestmates. Since it was easier to initiate emigra- tions rather than raids, most of the experiments were conducted during colony emigration. Several new exocrine glandular struc- tures have recently been discovered in ponerine ants (Holldobler and Haskins 1977; Holldobler and Engel 1978; Holldobler et al. 1982; Maschwitz and Schonegge 1977; Jessen et al. 1979). The Cerapachyini were not included in these studies. We therefore con- ducted first a histological survey for possible exocrine glands that might be involved in the communication behavior of Cerapach-)IS. Besides the known glands associated with the sting, we found a pygidial gland, which consists of a paired group of a few glandular cells under the 6th abdominal tergite. Each cell sends a duct through the intersegmental membrane between the 6th and 7th tergite (Fig. 9). The intersegmental membrane is laterally slightly invaginated, so that at each side it forms a small glandular reservoir. No particular cuticular structure on the pygidium is associated with the pygidial gland.
ln a first set of pilot experiments 1 dissected out of freshly killed Cerapach-I~S workers poison glands, Dufour's glands, hindguts, pygidial glands (6th and 7th tergites) and the last 3 sternites. For each test one organ of a kind was crushed on the tip of hardwood applicator sticks. These were then immediately inserted into the nest tube until the tip of the applicator was 2-3 cm away from the colony,



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Figure 9.
(a) Longitudinal section through the gaster of a Cerupwhss worker 5howing the location of the pygidial gland (PG), (b) Longitudinal section through the pygidial glank
GC: glandular cells; CH: glandular channels through inter- jegrnenta! membrane.




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18 Psyche [VOI. 89
which usually had gathered near the cotton plug. In the following 30 seconds 1 observed the reaction of the ants, and between each test 1 waited at least 10 minutes before another sample was inserted into the nest tube. These pilot tests (3 repetitions with each organ) clearly indicated that only crushed poison glands and pygidial glands eli- cited increased locomotory activity and attraction in Cerapa*s workers. The ants did not exhibit any particular behavioral reaction when sternites, hindgut or crushed Dufour's glands were intro- duced.* For the next series of experiments I first initiated colony emigrations either by following the procedure described above, or by shaking the colony out of the nest tube onto the arena floor. Before each experiment the arena was provided with a new paper floor. A new nest tube was offered 15-20cm away from the old nest tube or the displaced colony.
Once the colony emigration to the new nest tube had commenced, I covered the floor area between the colony and the new nest site with a cardboard, onto which I had drawn two artificial trails, one with a crushed glandular organ to be tested, and a second one with a drop of water (control). The trails were made to originate either from the entrance of the nest tube or from the periphery of the clustered colony. Each trail (test and control) diverged through an angle of 45O to either side from a possible natural trail (which was of course covered by a piece of cardboard). In addition the whole paper floor was rotated for 90å¡ in order to control for possible visual orientation (Fig. 10). During the following 2 minutes 1 counted the ants following the trails (10cm long) to the end. Only trails drawn with crushed poison glands elicited a precise trail fol- lowing behavior in Cerapachys workers. There was some initial following response to trails drawn with crushed pygidial glands, but the ants followed only through the first 1-3 cm, then usually turned or meandered off the trail. Only once was it possible to conduct a similar test during raiding behavior of Cerapachvs. In this instance the ants followed only an artificial trail drawn with a crushed poison gland.
Although pygidial gland secretions did not release trail following behavior in Cerapachys, it clearly elicited increased locomotory *Cerapachys has also a very well developed sting sheath gland. It was not possible to test whether or not secretions of the gland play a role in communication.



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19821 Holldobler - Cerapachys 19
NII
Figure 10.
Schematical illustration of the experimental arrangement during trail tests. The colony was emigrating from nest N1 to nest N11 along a natural trail a. During the trail tests, the whole arrangement was turned 90å (arrow). The natural trail a was covered by a cardboard, on which the test trail (T) and a control trail (C) were offered, each deviating from a in an angle of 45O. activity and attraction in the ants. I hypothesized therefore that the recruiting ant might discharge pygidial gland secretions when it exhibited the gaster raising behavior. The pygidial gland pheromone might function as an additional recruitment signal by which the recruiting ant keeps the raiding party stimulated when leading it to the prey colony. In order to test this hypothesis, 1 tried on four different occasions to close the opening of the pygidial gland by applying collophonium wax between the 6th and 7th tergites. Unfor- tunately these experiments failed; apparently the ants were too dis- turbed by the procedure. During two raiding expeditions of



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20 Psyche [Val. 89
Cerapams we succeeded, however, in diverting individual ants from the raiding column over a distance of at least several centi- meters by presenting two applicators in front of them, one contami- nated with pygidial gland secretions and the other with water. Both applicators were slowly moved away from the columns in opposing directions. Of a total of 10 ants tested, 4 responded by following for a few centimeters behind the applicator with the pygidial gland secretions; no ant followed the control applicator. Although these results can be considered only preliminary, they do suggest that pygidial gland secretions might be involved in the recruitment pro- cess of Cerapachys. This suggestion was further supported by the results of a series of experiments in which I offered artificial trails drawn with crushed poison glands. I compared the trail following response of Cerapa*~ (within the first two minutes) successively either to trails drawn with poison gland secretions only or to poison gland trails offered simultaneously with pygidial gland secretions. For each kind a total of 6 experiments was carried out. Between each test at least one day had elapsed. The following response appeared to be stronger to poison gland trails when offered together with pygidial gland secretions (5.5 k 2.9) than to those offered with- out pygidial gland secretions (3.0 2 1.4) (0.1 > p > 0.05; Students t-test). Because of lack of material this series could not be extended, and thus the results remain only suggestive. The two final experiments demonstrated that a trail (10cm long) drawn with one crushed poison gland, was still effective as an orientation cue several hours after it had been drawn. Using the same experimental arrangement described above (Fig. lo), I was able to show that emigrating Cerapachys would follow poison gland trails, 2 and 6 hours old, when they were offered after the natural trail had been covered. On the other hand, crushed poison glands introduced into the nest tube after 2 and 6 hours, or poison gland trails offered 2 and 6 hours after they had been drawn, did not elicit excitement or spontaneous trail following behavior. From these results it appears that the poison gland material might contain a short lasting stimulating component as well as a longer lasting orienting component.




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Holldobler - Cerapachys
Raiding expeditions in Cerapachys turneri are organized by indi- vidual scout ants, that return to the colony after having discovered a nest of the prey species. The scout lays a chemical trail with secre- tions from the poison gland, which serve as recruitment and orienta- tion signals for the nestmates. Circumstantial evidence suggests that in addition the scout releases a stimulating chemical recruitment signal from the pygidial gland. This occurs probably when the scouts move with their gaster held slightly upwards in a calling position.
Wilson (1958) reports the field notes made by H. Potter on the cerapachyine species Phyracaces potteri, which contain the only available description of the early stages of a complete raid observed in the field. Before the raid started Potter noted a few workers moving rapidly about, "each with its abdomen raised upwards". These observations match closely my findings in the laboratory and lend further support to the hypothesis that in addition to the trails laid with poison gland secretions, another stimulating signal is dis- charged, presumably from the pygidial gland of the recruiting ants. Wilson (1958) observed groups of Phyracaces moving along a raiding trail laid down by a raiding party on the previous day. In this case no individual leadership was involved and the foragers seemed to emerge from the nest randomly without a special recruit- ment stimulation by scout ants. Obviously these ants were following an established foraging trail, leading to a previously raided Pheidole nest which appeared to be vacated this time. Small exploratory parties conducted brief excursions to the side, but in most cases they turned back to the main trail. No nest suitable for raiding was found during these explorations.
These observations strongly suggest that chemical trails laid dur- ing raiding expeditions might still function as orientation cues one day later and that foraging parties can follow these established trails without the leadership of a recruiting scout ant. Indeed, my labora- tory experiments with Cerapachys have demonstrated that artificial trails drawn with poison gland material are effective as orientation cues at least for several hours.




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22 Psyche [vo~. 89
Although the raiding cerapachyine ants are usually enormously outnumbered by the worker force of the prey species, not one Cera- pachys worker was lost during all the raiding experiments in the laboratory. As can be seen from Fig. 2, Cerapachys and Sphinc- tomyrmex are excellently protected by a heavily sclerotized cuticle. The intersegmental joints, that is, the joints between head and thorax, and between thorax, petiole and gaster, are covered by cuticular projections so that no intersegmental membrane is ex- posed, even if the ant is twisted and bent to an extreme degree. In addition, Cerapachys and probably all the other cerapachyine ants have a most powerful sting that immobilizes the opponents within seconds. Not only the adults of the raided colony, but also the captured larvae and pupae are stung by the raiders before they are retrieved to the Cerapachys nest. Observations and experiments demonstrated that the prey larvae are kept in a stage of metabolic stasis and can thereby be stored for a period of more than two months. This food storage system enables Cerapachys to adjust the raiding activities to food requirement and supply. From the labor- atory experiments we can conclude that Cerapachys does not con- duct daily or periodic raiding expeditions. The frequency of raiding expeditions depends on the food supply stored inside the Cera- pachys nest.
I was unable to demonstrate periodic nomadic behavior in Cera- pachys in the laboratory. 1 assume that nest emigrations might occur relatively frequently in this species, but that they do not fol- low a periodic pattern. Instead, environmental factors such as food supply or physical conditions of the nest site are likely to play the important role in inducing a Cerapachys colony to emigrate. Many thanks to H. Engel-Siege1 for technical assistance, to E. Seling for the SEM work, and to W. L. Brown and R. W. Taylor for identifying the ants. I am most grateful to R. W. Taylor and the Division of Entomology, CSIRO, Canberra (Australia) for their generous hospitality. This work was supported by a grant from the National Science Foundation BNS 80-021 61 3, the National Geo- graphic Society and a fellowship from the John Simon Guggenheim Foundation.




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19821 Holldobler - Cerapachys
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A new sternal gland in ants and its function in chemical communicatio n . Naturwissenschaften in press.
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Neue Abdominaldrusen bei Ameisen. I. Ponerini (Formicidae: Poner-i- nae). Zoomorphologie 94, 49-66.
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1962. Das Paralysierungsproblem beim Bienenwolf, Philanthus iriangulum F. (Hym. Sphec.) Z. Vergl. Physiol. 45,413-462. RATHMAYER, W.
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Venoms of Sphecidae, Pompilidae, Mutilidae, and Bethylidae. Hand- book of Experimental Pharmacology vol. 48, Arthropod Venoms ( S - Bettini, ed.) pp. 661 -690. Springer-Verlag, Heidelberg-New York, 197 8 - WHEELER, G. C.
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Ant larvae of the subfamily Cerapachyinae. Psyche 57, 102-1 13. WHEELER, G. C. AND J. WHEELER
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The ant larvae of the subfamily Cerapachinae. Suppl. Proc. Entomol. Soc. Washington 66, 65-7 1.
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The Insect Societies. Belknap Press of Harvard University Press. Cam- bridge (Mass.).




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