Behav Ecol Sociobiol (1997) 40: 337 ± 349 Ó Springer-Verlag 1997

Bertrand Schatz á Jean-Paul Lachaud á Guy Beugnon

Graded recruitment and hunting strategies linked to prey weight

and size in the ponerine ant Ectatomma ruidum

Received: 27 June 1996 / Accepted after revision: 3 March 1997

Abstract According to the weight and size of their prey, played by E. ruidum workers is discussed in relation to
Ectatomma ruidum workers can employ di€erent re- their involvement in scavenging or predatory behavior.
cruitment systems (solitary hunting, cooperative hunting
and group hunting with recruitment) when mastering Key words Predation á Behavioral ¯exibility á
and retrieving prey items from short distances from the Graded recruitment á Ponerine ants á Ectatomma
nest. Prey size determined the backwards entry typically
adopted by this species, while prey weight determined
the predatory strategy selected. After a common initial
sequence (search for prey, detection, localization), pre- Introduction
datory sequences varied in terms of the type of ap-
proach, the site of seizure, the reaction after stinging and For all animals, regardless of whether they are solitary
the type of transport. Nevertheless, irrespective of prey or social, the search for and retrieval of food items
weight and size, seizure was preferentially oriented to- constitute the major part of their time-budget and en-
wards the head and prey were always stung. Short-range ergy expenditure. However, in social vertebrates, such as
recruitment and mass recruitment without trail laying lions, wolves or hyenas (Curio 1976; Alcock 1993), as
were elicited by a large range of heavy prey (> 2.5 times well as in social insects (Wilson 1971; Oster and Wilson
the weight of an individual worker). According to the 1978; Franks 1986; Mo€ett 1988; Schmid-Hempel 1991;
mortality risk associated with each prey, hunters ex- Duncan and Crewe 1993), cooperative food source
hibited a ``prudent'' stinging posture associated with an mastering and/or recovery may greatly enhance the ef-
increase in the duration of the subsequent phase of ®ciency of the foraging by reducing total energy costs.
waiting for prey immobilization. The overall time of In the course of their evolution, ants have developed
capture was positively correlated with the weight of the a variety of foraging strategies depending on their spe-
prey. When collective hunting strategies were involved, cies-speci®c behavioral repertoire and communication
E. ruidum colonies matched the number of recruited systems, and on various parameters related to the food
hunters to the size and weight of the prey. Compared to sources (e.g., nature, size, weight, quality, availability,
solitary hunting strategies, for short food±nest distances, accessibility) and to the intrinsic characteristics of the
this graded recruitment appeared to enhance the ener- foragers (e.g., size and strength, level of internal ener-
getic bene®ts derived by this species from the use of getic reserves, prior experience, nutritional status of the
recruitment systems: the higher the number of workers colony) (Traniello 1989a; HoÈlldobler and Wilson 1990;
involved in the recruitment process, the greater the en- Schmid-Hempel 1991). From an adaptive perspective,
ergetic bene®ts obtained. The exhibition or absence of the more ¯exible the foraging behavior, the more readily
trail laying behavior in the recruitment responses dis- colonies may adjust to environmental changes. Bernstein
(1975) found that the shifts in foraging strategies re-
corded in the seed harvesting ants Pogonomyrmex spp.
B. Schatz á J.-P. Lachaud á G. Beugnon and Messor (ˆ Veromessor) pergandei, were related to
Laboratoire d'Ethologie et Psychologie Animale, seed density. At high densities, these species exhibited
U.M.R.-C.N.R.S. 5550, Universite Paul-Sabatier,
118 Route de Narbonne, F-31062 Toulouse Cedex, France individual foraging. At medium densities, scout ants
were used to locate food and to recruit a column of ants.
J.-P. Lachaud (&)
El Colegio de la Frontera Sur,
At the lowest densities, group foraging without a scout
Apdo. Postal 36, 30700 Tapachula, Chiapas, MeÂxico ant occurred along de®ned trunk trails. A similar rela-
Fax: (52) 962-81015 tion between the relative food abundance and the for-
338

aging strategy adopted by the large African ponerine ant conditions under which they are induced, we studied the
Megaponera foetens, was proposed by Longhurst and ¯exibility of the workers' predatory behavior when faced
Howse (1979) to account for the regional di€erences in with prey of di€erent sizes and weights. These factors, as
foraging and recruitment systems reported for this well as the distance of the food source from the nest,
termitophagous species. For a given species, the exis- appear to be the key parameters responsible for the
tence of a multiple strategy of foraging is mainly at- energy cost related to food retrieving and determining
tributable to its ability to recruit nestmates to food the choice of predatory strategy, especially in species
sources and to the variety of the recruitment systems generally considered as solitary foragers (see Dejean
employed (Wilson 1971; Oster and Wilson 1978; HoÈll- 1982, 1991; Dejean et al. 1993; Lachaud and Dejean
dobler and Wilson 1990). Such ¯exibility in foraging 1994).
techniques may permit the utilization of alternative food
sources by specialized species (Dejean 1982; Masuko
1984; Lachaud and Dejean 1994) or the exploitation of a
wider range of food items by generalist species (HoÈll- Methods
dobler 1984; Dejean et al. 1993).
The ponerine ant Ectatomma ruidum Roger is one of Five queenright colonies of E. ruidum, ranging from 70 to 350
these generalist species. This terricolous medium sized workers, were reared in plaster nests placed in an experimental
room under controlled conditions (temperature at 25 °C ‹ 1 °C,
ant (7±9 mm in length; 5±12 mg in weight, mean = 60% ‹ 5% humidity and a 12/12 L/D photoperiod). For each
8 mg) is widespread in disturbed habitats as well as in colony, a 35 ´ 35 cm box, connected to the nest, was used as a
damp rain-forests (Brown 1958). Very common in foraging area. Ants had access to a permanent supply of honey and
Central and northern South America (Weber 1946; water, in addition to fresh prey (crickets, cockroaches, or Tenebrio
Kugler and Brown 1982; Levings and Franks 1982; larvae) given twice a week. In this study, we examined in laboratory
conditions the predation sequence exhibited by the hunters when
Young 1986), it is the dominant ant species in some o€ered larvae of Tenebrio molitor as prey. In order to avoid pre-
habitats like tropical thorn forest in Colombia (Kugler datory specialization or preference associated with a particular type
and Hincapie 1983) or co€ee and cocoa plantations in of prey, instead of several types of prey with di€erent sizes, we
southeastern Mexico, where its density can vary between chose to use only one type of prey showing large di€erences in size.
Prey (n = 274) di€ering in size and weight (respectively 7±27 mm
2 700 and 11 200 nests per hectare (Lachaud et al. 1996). in length, 4±204 mg in weight) were o€ered randomly. Prior to each
Its eciency as a predator and its ecological impact on experiment, the prey o€ered was measured and weighted, and the
insect pests had already been noted and various authors observation of the predation sequence began when the prey was
have proposed this species as a potential agent for bio- placed at the center of the foraging area (about 20 cm from the nest
entrance). A signi®cant positive logarithmic correlation was de-
logical control in co€ee/cocoa plantations in Panama tected between prey size and weight (r = 0.977; P £ 0.0001).
and Mexico (Weber 1946; Lachaud 1990), and in maize The duration of predation tests was ®xed at 30 min for all ex-
cultures in Nicaragua (Perfecto 1990, 1991). Whereas a periments. In all cases, the exact behavioral sequence and its du-
substantial body of information exists concerning vari- ration were recorded. A squared paper, placed under the trans-
ous aspects of its foraging activity, social behavior and parent ¯oor of the foraging arena, allowed us to estimate the dis-
tance of detection of the prey (as determined by a change in the
intraspeci®c relationships (Lachaud et al. 1984; Ja€e and motility and orientation of the ants to the prey, see Dejean 1982).
Marquez 1987; Lachaud and Fresneau 1987; Corbara The orientation of the hunter towards the prey during the approach
et al. 1989; Pratt 1989; Breed et al. 1990, 1992; Lachaud and the part of the body prey seized during the attack were also
1990; Schatz et al. 1994), little information is available analyzed. In the case of strategies for which several hunters had
been involved, only the behavior of the ®rst hunter was considered.
on its predatory behavior. In the laboratory, foragers of In order to discriminate between weight and size e€ects, the
E. ruidum were observed to fall into one of ®ve spe- weight of 44 additional larvae of T. molitor, belonging to a given
cialized categories: hunters, sugar collectors, unspecial- size category (12.5±14 mm in length), was reduced by removing a
ized intermediates, patrollers, and nest-maintenance certain amount of their haemolymph with a syringe needle. Im-
mediately after the wounded larvae ceased to bleed, it was placed in
workers (Schatz et al. 1995). There is an even more the foraging area. Treated larvae were alive and moving at the
subtle within-caste specialization between stingers and moment of attack by E. ruidum workers.
transporters within the hunters (Schatz et al. 1996). Observations were sorted a posteriori, according to the strategy
While this species has been described as a solitary employed by the ants towards prey of di€erent weight and size. For
hunting ant (Levings and Franks 1982; Lachaud et al. each predatory strategy, a ¯ow diagram was devised from obser-
vational data. For each sequence, percentages were calculated
1984), recruitment may occur in some instances (Lac- based on the overall number of transitions between each individual
haud 1985; Pratt 1989) and some degree of cooperation behavioral act. For strategies involving recruitment, two parts were
between stingers and transporters has been reported considered for the calculation of the transition frequency between
(Schatz et al. 1996). However, the available information each behavioral act: (i) one hunter only, and (ii) all hunters par-
ticipating in the predatory sequence. During cooperative hunting,
dealing with recruitment behavior appears somewhat recruited individuals were involved in di€erent steps of this se-
contradictory (see Lachaud 1985; Pratt 1989; Bestmann quence. Therefore in the calculation of transition frequencies, we
et al. 1995), and the factors which provoke the exhibi- considered the total number of individuals starting at the beginning
tion of trail-laying behavior or its absence remain un- of the recruitment phase.
Complementary ®eld observations were made at the beginning
clear. of the rainy season, in the same co€ee plantation from where the
To determine the predatory repertoire of this species colonies studied in the laboratory had been collected. This plan-
and to analyze the cooperative strategies used and the tation was located in the Rosario Izapa Experimental Station of the
 339

Instituto Nacional de Investigaciones Forestales y Agropecuarias, prey weight was 8 mg (10±90% range: 4.9±11 mg), and
in the Cacahuatan District, Chiapas State, Mexico, at 420±440 m for type 2, the median weight was 15 mg (10.5±19.5 mg).
elevation. In order to estimate cooperative foraging in this species,
six nests of E. ruidum were observed over 6 h every day, for 4 The di€erences in median weight and range of prey
consecutive days. Observations were made between 0830 and 1430 weights for each of the three remaining strategies were
hours, a period which includes nearly 60% of the daily activity in much greater: 39 mg (17.6±79.0 mg) for the cooperative
the rainy season (Lachaud 1990). We recorded the type and the hunting, 115 mg (72±157 mg) for type 1 group hunting
number of prey brought back to the nest by solitary ants or groups
working collectively. The range and distribution of prey weights
with recruitment, and 175 mg (159±199 mg) for type 2
and sizes was estimated by measuring a sample of prey (earth- group hunting with recruitment. In the latter case,
worms) showing cylindrical-shape analogies with the T. molitor however, the possibility of a more extensive range of
larvae used in the laboratory experiments. prey weights could not be ruled out since observational
data were limited by the size/weight characteristics of the
experimental prey.
Results

Three main types of predatory strategy were de®ned Common behavioral phases
depending on prey weight and size: solitary hunting,
cooperative hunting involving only short-range recruit- In this study, as far as the succession of the main
ment, and group hunting with recruitment from the nest. behavioral phases is concerned, the ®rst part of the
On the basis of qualitative changes in the behavioral behavioral sequence (search for prey ± detection/
sequence, it was possible to distinguish some variants localization ± approach/antennation ± seizure) was
both in the solitary hunting strategy (according to the essentially the same for all the strategies. Nevertheless,
frequency of the lifting, stinging and ``let go and re- some aspects can vary, namely those related to the ori-
seize'' phases, and to the mode of entry into the nest) entation of hunters towards the prey and the part of the
and in the recruitment strategy (according to the number body prey seized.
of ``givings-up'' and the whether or not the prey was
cut). However, in all cases, a certain ¯exibility was en-
countered in the choice of the strategy adopted. The Search for prey ± detection/localization
range of prey weights corresponding to each strategy
was not strict: two prey items with the same weight During the search for prey, the worker moved slowly
could trigger di€erent responses. For this reason, the and sinuously, antennae spread apart and mandibles
median weights and the 10±90% ranges were determined closed. For all cases, detection occurred as soon as the
for each strategy. prey appeared to enter the ®eld of visual perception of
Solitary hunting involved a small range of prey the ant. Immediately, the hunter oriented its antennae
weights (Fig. 1). For solitary hunting type 1, the median towards the prey and opened its mandibles. Prey were
detected at a distance of 1±2 cm, a ®gure similar to that
found in the ®eld (Lachaud 1990). However, the move-
ment of the prey was apparently necessary for its de-
tection and localization, since various workers passed by
a motionless prey at this same distance of 1±2 cm
without locating it.

Approach/antennation ± Seizure

The solitary strategy type 1 di€ered from all other

strategies: it was the only situation for which no pref-
erential direction was detected during the approach, al-
though hunters clearly chose to seize the prey in the
middle part of its body. For all other strategies, during
the approach, the middle part of the body of the prey
was signi®cantly preferred, but the head was also chosen
preferentially to the tail. Moreover, for the site of the
Fig. 1 Hunting strategy adopted by Ectatomma ruidum workers seizure, a highly signi®cant preference was shown for the
according to prey weight. Whisker box representation with median head or, to a lesser extent, for the tail; hunters avoided
weight of the Tenebrio molitor larvae o€ered as prey (middle bar) and the central part of the body prey (Fig. 2). During the
20±80% range values (extremities of the whisker box); open circles seizure phase, the hunter displayed a forward movement
indicate the observed minimal and maximal values; bars indicate the
10±90% range values. There is some overlap at the division between
of the whole body. With the exception of the solitary
adjacent categories of prey weights: two pieces of prey with the same strategy type 1, the seizure behavior normally occurred
weight can trigger two di€erent predatory responses according to the following sequence. When its antennae
340

Fig. 2 Part of prey body used as target during approach and seizure **0.001 < P < 0.01, ***0.0001 < P < 0.001; ****P < 0.00001;
phases for the di€erent hunting strategies. Chi-square comparison n = number of individual approaches or seizures)
with a theoretical random approach or seizure (*0.01 < P < 0.05,

came in contact with a motionless prey, the hunter po- Solitary hunting
sitioned its body above that of the prey in order to align
its body axis with that of the prey, and always oriented Two variants of solitary hunting sequences, related to an
its head towards the head of the prey. Once this posture increase in the weight and size of the prey, were distin-
was secured, the hunter opened widely its mandibles, guished: type 1 and 2 (Fig. 3a, b). Solitary hunting type 1
placed its antennae on each side of the body of the prey, occurred with the smallest prey, ranging from 4 to 12 mg
and went up until the head where the seizure abruptly in weight and 7 to 12 mm in length. The total time of
occurred. This particular behavior strongly suggested capture lasted from 10 to 160 s, with a mean time of
that the preference exhibited by the hunter to seize its 77.6 ‹ 40.5 s (n = 44). Solitary hunting type 2 oc-
prey by the head, really corresponded to an active curred with prey ranging from 9 to 30 mg in weight and
choice. This sequence was the most common (121 times 11 to 17 mm in length. The total time of capture lasted
out of 204). However, other sequences were also ob- from 40 to 220 s, with a mean time of 141.2 ‹ 40.5 s
served. For instance, various hunters were observed (32 (n = 40). For both types, most abandoned prey (77%,
times) running over the body of the prey while walking n = 13) were encountered during the ``let go and re-
by its side and, when several hunters were involved in seize'' phase. When the prey did not struggle after being
the attack, while the ®rst hunter seized the prey by the stung, it was directly transported to the nest. If the prey
head, the second hunter seized it by the middle part of its struggled, the hunter stung it repeatedly and waited for
body (2 cases) or by the tail (16 cases) (comparison with its immobilization for at least 20 s before transporting it
a random seizure: v2 = 19.6; P £ 0.0001). This suggests to the nest.
that hunters prefer to seize the prey by their extremities Di€erences between both types were qualitative and
and more particularly by the head. quantitative. Aside from the duration of the total se-
 341

Fig. 3 Flow diagrams for the solitary hunting strategy: a solitary based on the overall number of transitions between each individual
hunting type 1 (n = 44 sequences); b solitary hunting type 2 (n = 40 behavioral act
sequences). For each sequence, percentage values were calculated

quence, the main di€erences concerned the lifting and weight of the prey, or solitary hunting type 2 if they were
stinging phases, of which the occurrence was less fre- more sensitive to the size of the prey. The predatory
quent for solitary hunting type 2, and the ``let go and re- strategy adopted by hunters faced with experimentally
seize'' phase which was much more frequent in this case. modi®ed prey corresponded to solitary hunting type 1,
Besides, while for solitary hunting type 1 the entry into except for entry into the nest (Table 1). In all cases, this
the plastic tube connected to the nest was always direct, latter behavior was performed backwards as in solitary
for type 2, after several attempts to enter directly, the hunting type 2. These data strongly suggest that prey
hunter always turned round and entered backwards, weight determines the solitary hunting strategy for
pulling its load. E. ruidum, except for the way in which the hunters enter
In order to distinguish between the e€ects of the size the nest, which is determined by the size of the prey item
of the prey and its weight in the choice of the solitary (probably for mechanical reasons, Lachaud 1985).
strategy to adopt, we experimentally manipulated the
weight of similar size prey (see Methods). A total of 44
larvae of T. molitor were chosen within a given range of Cooperative hunting
intermediate sizes (between 12.5 and 14 mm) corre-
sponding to a median of 15 mg (range of weights: 13± Retrieval of heavier and larger prey (approximately
17 mg), for which hunters regularly adopted solitary ³ 20 mg in weight and 15 mm in length) appeared to be
hunting type 2. By extraction of haemolymph, prey dicult for a solitary individual. In these cases, the co-
weight was reduced to a median of 8.5 mg (range of operation of several workers was generally necessary
weights: 7.5±9.25 mg), for which hunters normally and was triggered by the scout ant.
adopted solitary hunting type 1. Faced with these ex- Cooperative hunting (Fig. 4) occurred for prey of a
perimental prey, we predicted that hunters would adopt range of weights and sizes from 13 to 98 mg and 13 to
solitary strategy type 1 if they were more sensitive to the 22.5 mm, respectively. The total time of capture lasted
342

Table 1 Comparative importance of size against weight of prey in from about 15 mg to 8.5 mg by removal of haemolymph. Prey size
the choice of predatory strategy to adopt by solitary hunters of determined the backwards entry into the nest while prey weight
Ectatomma ruidum. Treated prey refer to 44 larvae of Tenebrio determined the choice of predatory strategy (chi-square compar-
molitor (12.5±14 mm in length) in which mean weight was reduced isons)

Behavioral acts Number of behavioral acts performed Statistical comparison (v2 test)

With treated During solitary hunting During the solitary Between Between
prey (n = 44) type 1 (n = 44) hunting type 2 (n = 40) A±B A±C
(A) (B) (C)

Lifting of prey 42 42 24 NS v2 = 15.6

P = 0.0001
Direct transport of 18 16 1 NS v2 = 17.7
motionless prey P < 0.0001
Leaving moving 9 10 27 NS v2 = 18.9
prey P = 0.0001
Stinging moving prey 15 16 11 NS NS
Backwards entry 38 0 38 v2 = 66.9 NS
into the nest P < 0.0001

from 80 to 990 s, with a mean time of capture of strategy were never recruited directly from the nest by
418.2 ‹ 273.3 s (n = 39). By contrast with both strat- the scout ant, but were attracted from the surroundings
egies of group hunting with recruitment described be- of the prey. The scout ant, after the ®rst attempt at
low, the recruited workers involved in the cooperative seizure, engaged in repeated, aggressive stinging. The

Fig. 4 Flow diagram for the cooperative hunting strategy (n = 40 sequences)

 343

nearby nestmates (within a radius of about 10 cm) ap- discovered, the typical succession of behavioral acts
peared to be attracted and readily joined the scout ant (detection, localization, approach, antennation, seizure,
and began to attack the prey and sting it. Although no stinging, and dropping) was observed. Heavy prey were
evidence of any chemical communication was detected, never lifted. A rudimentary mass recruitment without
the release of an alarm pheromone originating from the trail laying was then triggered by the scout ant which
mandibular glands or the poison gland (Pratt 1989) may performed a very excited return path back to the nest in
have occurred. Visual and acoustical information could order to activate other hunters prior to returning to the
also be responsible for the attraction observed. prey (see Lachaud 1985; Pratt 1989). The orientation of
Of a total of 39 prey which triggered local recruit- the recruited ants towards the prey was not related either
ment, all were successfully transported to the nest. to the presence of a leader nor with a chemical trail, but
Nevertheless, only 71.1% of the hunters (of a total of appeared to be directed by signals originating from the
121 involved in the cooperative hunting) participated in prey (odors, movements, visual or acoustic informa-
all the phases of the killing and transport of the prey to tion). Evidence for this was supported by the fact that
the nest. each recruited ant followed a di€erent individual path
between the nest and the position of the prey which,
nevertheless, came together at the prey. Ants recruited
Group hunting with recruitment by the scout ant never engaged in recruitment behavior.
Subsequently to this ®rst recruitment phase, two
Two variants of group hunting with recruitment were di€erent behavioral sequences were exhibited according
distinguished following an increase in prey weight and to the weight of the prey (Fig. 5a, b). Group hunting
size (for prey approximately ³ 79 mg and 21.5 mm). with recruitment type 1 occurred with prey ranging from
Both were characterized by a similar behavioral se- 61 to 165 mg in weight and 20.5 to 25.5 mm in length.
quence performed by the recruiter. Once the prey was The total time of capture lasted from 670 s to more than
30 min. Of a total of 39 prey which triggered this
Fig. 5 Flow diagrams for the group hunting strategy with recruitment:
strategy, 36 (92.3%) were successfully retrieved. Never-
a group hunting with recruitment type 1 (n = 42 sequences); b group theless, only 58.9% of the hunters (of a total of 214
hunting with recruitment type 2 (n = 108 sequences) involved in the recruitment) participated in all the
344

phases of the killing and transport to the nest. Group and 5): their hindlegs were raised high above the gaster
hunting with recruitment type 2 occurred with prey in an apparently defensive posture. The use of this
ranging from 152 to 201 mg in weight and 24.5 to ``prudent'' stinging by the hunter was clearly associated
27 mm in length. The upper limits of 201 mg and 27 mm with an increase in the duration of the subsequent phase
resulted from the limits in the weights and sizes of of waiting for prey immobilization (from 20±40 s to
available T. molitor larvae in our studies. Nevertheless, more than 150 s). A total of 126 cases (out of 420
the great increase in both the number of giving-up be- stinging behaviors) were recorded over the three collec-
haviors before recruitment (85.2%) and the total dura- tive strategies (23.1%, 29.5%, and 32.2% of the sting-
tion of the capture (which in all cases lasted more than ings, respectively), for which the ``prudent'' stinging
30 min) suggested that a prey weight of 200 mg actually posture was exhibited by the hunters: a subsequent
constituted a limit for the hunters of E. ruidum to phase of extended wait for prey immobilization occurred
complete the predatory sequence. The main character- in 83.3% of these cases, while such an extended wait was
istic of this sequence was the frequent cutting of the prey never observed after ``normal'' stinging. Moreover, the
(28.1%), always performed between two segments of the increase in the percentage of ``prudent'' stinging fol-
Tenebrio larva. A total amount of 162 individuals was lowed by an extended wait for prey immobilization ac-
recruited, i.e. an average of 10.1 recruited ants per re- cording to the increase in the weight of the prey (44.4%
cruiting worker, signi®cantly di€erent (v2 = 6.87; for cooperative hunting, 82.3% for group hunting with
P £ 0.01) from the average of 4.5 recruited ants per re- recruitment type 1, and 90.9% for group hunting with
cruiting worker reached in the previous type of group recruitment type 2) suggested that hunters adjusted their
hunting with recruitment. Of 16 prey which triggered behavioral response, perhaps related to a ``mortality
group hunting with recruitment type 2, 13 (81.3%) were risk'' associated with each prey.
successfully brought back to the nest. Only 46.1% of the
recruited hunters (of a total of 178 hunters involved in
the recruitment) participated in all the phases of the Duration of capture and number of hunters involved
killing and transport to the nest.
The time needed to capture a prey item and transport it
to the nest was positively correlated with its weight
(r = 0.952; P < 0.0001; n = 140) (Fig. 6a). The hunt-
Comparative analysis of some characteristics
ers demonstrated an important ability to adjust the
number of recruited nestmates according to the weight
Lifting of the prey and stinging behavior
of the prey. In spite of the variability in the defense
behavior performed by the prey triggering the di€erent
In both types of solitary hunting, the small and light
strategies, a highly signi®cant positive correlation was en-
prey were lifted, in most cases, and rapidly stung by the
countered between prey weight and number of hunters in-
hunters. Nevertheless, in the subsequent collective
volved in the predatory sequence (r = 0.962; P < 0.0001;
strategies, this behavior rapidly disappeared with the
n = 94) (Fig. 6b). Moreover, this positive correlation
increase of the weight and size of the prey. When the
persisted within each one of the collective strategies: the
prey were larger and heavier (>15 mm in length and
number of hunters involved varied from 2 to 8 individ-
20 mg in weight), hunters of E. ruidum began to adopt a
uals for cooperative hunting (r = 0.821; P < 0.001;
particular posture during the stinging behavior (Figs. 4
n = 39), from 6 to 11 individuals for the group hunting
Fig. 6 Relationship between prey weight and a duration of the
with recruitment type 1 (r = 0.522; P < 0.05; n = 39),
predatory sequence or b number of workers involved in the predatory and from 8 to 15 individuals for the group hunting with
sequence recruitment type 2 (r = 0.686; P < 0.01; n = 16).
 345

Eciency of the predatory strategy adopted all the hunting strategies described above. While di€er-
ent types of prey were involved, the ®ve strategies ob-
According to the prey weight, the predatory eciency of served in laboratory studies with larvae of Tenebrio
E. ruidum, as measured by the ratio of actual captures molitor could be easily extrapolated to the diversity of
per hunting attempt was relatively high (84.1±92.3%) natural prey. However, the frequency of collective prey
and only the very large prey (weight between 20 and 25 recovery was very low: 96.4% of prey were brought back
times that of an individual ant) induced frequent giving- to the nest by solitary ants. Moreover, the range and
up by the hunters (eciency of retrieval 12%), which distribution of the weights and sizes of 84 earthworms
occurred essentially during the solitary phase prior to (Halotaxida, Lombricidae) separated from the collected
any recruitment. In the case of the collective hunting prey, showed that though 15.5% of this type of prey
techniques, however, considering only the attempts were larger than 20 mm in length, only 1.2% (1 case)
which actually triggered recruitment behavior, it is ob- was heavier than 20 mg. This result con®rms that the
vious that all three collective strategies used according to range of weights and sizes of the prey collected in ®eld
the increase in prey weight were highly ecient, with a conditions corresponds to that triggering the solitary
subdue-and-retrieval eciency varying from 81.3% to hunting strategy in laboratory conditions.
100%.
This eciency of the collective hunting techniques
was accompanied by an increase in the net energetic
bene®t derived from the cooperative prey retrieval. Discussion
Considering the mean number of workers involved in
prey retrieval and the mean weight of prey retrieved Behavioral sequences
successfully to the nest, the overall mean energetic ben-
e®t (Table 2) was greater for collective strategies than Although, in ®eld conditions, E. ruidum workers appear
for solitary ones, given a distance of only 20 cm from to forage individually for the most part, they can employ
nest entrance. Furthermore, if we consider that only a di€erent recruitment systems when mastering and re-
part of the total workers recruited in each one of the trieving large prey items. The behavioral analysis of
collective strategies actually completed all the steps of these infrequent collective retrieval strategies appears
the predatory behavior involved in the transport of large worthwhile considering the possibly strong adaptive
prey to the nest, it may be argued that the coarse esti- value of such ¯exibility in the predatory behavior of this
mates of energetic bene®ts indicated here are underesti- species. According to the increasing weight and size of
mates of the actual bene®ts derived from the use of the prey, ®ve sequences of predatory behavior, clustered
recruitment systems by E. ruidum. Overall, these esti- into three main strategies, were distinguished: solitary
mates suggest that the mean weight of prey retrieved per hunting, with two variants, for small prey (weight < 2.5
individual worker actually increased according to the times that of an individual worker), cooperative hunting
mean number of hunters recruited by the scout ant. for medium prey (weight between 2.5 and 9 times that of
the ant) and group hunting with recruitment, also with
two variants, for large prey (weight > 9 times that of
Field observations the ant).
For predatory species, which are not equipped with
During the 24 h of observation dedicated to each nest, long mandibles, as is the case for E. ruidum, the ®rst
419 prey were collected belonging to 50 families and 19 behavioral phases of the predatory sequence are always
orders of arthropods (insects, arachnids, crustaceans, very short and consist in a rapid approach and anten-
diplopods, chilopods), annelids (earthworms) and mol- nation, followed by the seizure and the stinging of the
lusks (snails, slugs). During this survey of predatory prey (Dejean and Bashingwa 1985). In E. ruidum,
hunting in ®eld conditions, E. ruidum hunters exhibited whatever the size of the prey, the initial sequence was

Table 2 Estimation of the energetic bene®ts derived by E. ruidum estimate (n) was either the total number of workers involved in all
from the use of ®ve di€erent hunting strategies (solitary hunting cases corresponding to each strategy (minimum estimate) or only
type 1 and 2, cooperative hunting, collective hunting with recruit- the number of workers involved in all successful prey retrieving
ment type 1 and 2). The number of workers considered for this (maximum estimate)

Median prey weight Total weight Energetic bene®ts (in mg of retrieved prey per individual)
triggering each strategy retrieved
(mg) (mg) Minimum estimates Maximum estimates

8 284 6.45 (n = 44) 7.68 (n = 37)

15 515 12.88 (n = 40) 15.15 (n = 34)
39 1465 12.01 (n = 122) 17.03 (n = 86)
115 4102 18.90 (n = 217) 32.56 (n = 126)
175 2196 8.13 (n = 270) 26.78 (n = 82)
346

always observed according to the same succession of cerned, the duration of the waiting phase is signi®cantly
behavioral acts: search for prey, detection and localiza- longer when it occurs after the stinging behavior with
tion. The position of the antennae during this ultimate hindlegs raised. A similar behavior has been described
phase, suggests that olfaction could be important during for Pachycondyla villosa (Dejean et al. 1990) and Pal-
the localization of the prey (Dejean and Bashingwa tothyreus tarsatus (Dejean et al. 1993). According to
1985). Moreover, in E. ruidum visual information is these authors, it corresponds to a ``prudent'' posture, in
likely to be in¯uential since, as for Myrmecia nigriceps order to avoid the defensive kicks of the prey which may
(Via 1977; Eriksson 1985) and Odontomachus troglodytes injure the ant. This behavioral response suggests that,
(Dejean and Bashingwa 1985), prey movements appear when faced with vigorous large and heavy prey, scout
necessary to achieve localization by hunters. Such prey ants of E. ruidum are able to perceive the presence of a
movements are also likely to be involved in the short- danger and to associate a mortality risk to this kind of
range recruitment triggered by medium weight prey. prey. The perception of a mortality risk associated with
Considering the importance of visual information in the access to a food source has been demonstrated for
other aspects of its foraging behavior, like the orienta- some ant species (Nonacs and Dill 1988, 1990, 1991) and
tion towards food sources or the spatial specialization of was shown to a€ect their foraging patterns (Traniello
honey-collectors (see Ja€e et al. 1990; Schatz et al. 1994, 1989b; Nonacs 1990). In the case of E. ruidum, the fact
1995; Beugnon et al. 1996), the use of such information that large and heavy prey may be associated with a
in the predatory behavior of this species is not surpris- mortality risk seems to be con®rmed by the clear in-
ing. Finally, another factor, the nutritional state of the crease in the duration of the subsequent phase of waiting
hunter, is known to possibly a€ect the distance of de- for the immobilization of the prey (4±8 times greater).
tection of the prey (Holling 1966; Hardman and Turn-
bull 1980; Dejean 1987), and such a situation could not Recruitment systems and graded recruitment
be ruled out in our experiments. Nevertheless, food was
provided regularly to the colonies in small quantities and The factor that determined the strategy adopted by a
a possible e€ect of repletion or starvation was not con- scout ant of E. ruidum appeared to be prey weight. Prey
sidered as highly probable. size essentially a€ects the forward or backward entrance
During the approach, the hunter moves very slowly in of the hunter carrying its load, in a mechanistic way.
order to place itself in a favorable position for the at- Moreover, for the type of prey used in our study, taking
tack. Such an approach, very common in ants, is into account the positive logarithmic correlation between
thought to allow the hunter to avoid a subsequent attack size and weight, for large prey the range of variation of
from the prey (Dejean 1982). The high preference shown prey weights is much greater than that of prey sizes,
by the hunters of E. ruidum for the head of the prey as making prey weight a more useful parameter to estimate
site of seizure, is very similar to what occurs for other the energetic cost associated to a given prey item.
predatory ant species and, according to Dejean (1982), Two types of recruitment were involved during our
can be interpreted as an attempt to destroy the nervous experiments: a local, short-range recruitment which at-
system of the prey. tracted only the nearby nestmates and a rudimentary
According to the type of strategy, qualitatively im- mass recruitment performed in the nest without trail
portant changes occur in the behavioral sequence after laying. It is currently assumed (Sudd and Franks 1987)
the phase of the seizure of the prey. It may be argued that any recruitment system which reduces the number
that this phase is essential for the hunter to estimate the of ants returning without a load to their nest would be
size and weight of the prey and to determine the sub- an advantage. Nevertheless, the more the recruitment
sequent strategy to adopt. For prey which are not too system could adjust the response levels to additional
heavy (< 20 mg, i.e., with a weight < 2.5 times that of labor needs, the more ecient this recruitment would
the hunter) the vigorous struggle of the prey releases the prove to be (Taylor 1978; Oster and Wilson 1978). In
lifting and stinging behaviors of the hunter, as described some species, the reinforcement of the chemical trail
for Myrmecia gulosa (Robertson 1971), O. troglodytes between the food source and the nest is proportional to
(Dejean and Bashingwa 1985), Pachycondyla soror the food quality (i.e., the richer the food source, the
(Dejean 1991), Paltothyreus tarsatus (HoÈlldobler 1984; more pheromone is laid per returning forager), allowing
Dejean et al. 1993) and various dacetine ants (Dejean an adaptive response to food sources of di€ering quality
1982; Masuko 1984). Moreover, in E. ruidum, the rapid (Hangartner 1970; Verhaeghe 1982; Crawford and Ris-
®rst part of the predatory sequence (approach and an- sing 1983; Beckers et al. 1993; de Biseau and Pasteels
tennation) is always followed by the stinging of the prey, 1994). In a previous study on recruitment behavior in
as in Hypoponera sp., Ponera coarctata, Pachycondyla E. ruidum Pratt (1989) concluded that colonies did not
(ˆ Mesoponera) ca€raria, Aphaenogaster senilis and appear to match numbers of recruited workers to the
A. subterranea (Agbogba 1982). The stinging is always size of baits. On the contrary, our data demonstrate that
associated with a waiting phase of at least 20 s, except when collective strategies are used for the mastering and
for very small larvae of T. molitor which are immediately retrieving of large prey, and whatever the recruitment
transported to the nest by the hunter. In the case of the system involved (short-range or mass recruitment), the
collective strategies, and whatever the strategy con- number of nestmates recruited by the scout ant is posi-
 347

tively correlated with the weight of the prey. Further- di€erentiated maximal response. Moreover, in our ex-
more, for short-range recruitment, the number of ants periments, aside from being free to be moved, T. molitor
attracted may be progressively regulated in the course of larvae were vigorous living prey which were associated
the process by a possible feed-back mechanism: the with a mortality risk, which was unlikely to be the case
vigorous prey movements triggered by the stinging for a dead prey item. Such a di€erence might explain
(kicks, twists, starts) probably enhance the ability of why, for scavenging behavior, large prey trigger trail
workers to detect the prey. Moreover, as shown in laying even when the prey is located at an unusually long
Aphaenogaster (= Novomessor) albisetosus and cocker- distance (up to 7 m, see Pratt, 1989), while this form of
elli (Markl and HoÈlldobler 1978), the emission of recruitment is not performed for predatory behavior and
stridulations, known to occur mainly in the ultrasonic merely relayed by short-range recruitment or rudimen-
range in E. ruidum (Pavan et al. 1996), is likely to be tary mass recruitment by incitement to leave the nest.
involved as a modulatory signal (see HoÈlldobler 1995) Additional evidence suggesting that E. ruidum workers
enhancing the behavioral response of nestmates to this do not show the same behavioral response for scavenging
recruitment. For mass recruitment, however, the number and predatory behavior sequences is the di€erence ob-
of recruited ants has to be ®nely adjusted at the start served between the exploitation of either a source of 50±
since a unique foraging team (constituted of all recruited 100 freshly killed termites (Pratt 1989) or a source of 20±
workers) occurs in each case. Such a modulated response 100 living Drosophila ¯ies (Schatz et al. 1996). In the ®rst
of an individual hunter which, according to some char- case, when only scavenging behavior is concerned, the
acteristics of the food source, is able to determine scout ant returns to the nest to recruit nestmates laying a
whether to recruit and, if this recruitment occurs, the chemical trail while in the second case, dealing with
number of recruited foragers requested (Taylor 1978), predatory behavior, a collective strategy occurs with a
constitutes a typical example of graded recruitment. division of labor between the hunters, some individuals
The only other case of graded recruitment in a po- specializing as killers, the others as transporters. Thus,
nerine ant was reported by Breed et al. (1987) for an- the range of modulated behavioral responses available
other species of the Ectatommini tribe, the giant tropical for E. ruidum foragers, according to the situation, ap-
ant Paraponera clavata. Individual workers of P. clavata pears very impressive and the trail laying behavior de-
encountering prey items, gauge the size and/or un- scribed by Pratt should in fact correspond to the highest
wieldiness of the item regardless of its weight when de- level in the graded series of mass recruitment used by this
termining whether to recruit. Nevertheless, as pointed species.
out by these authors, the prey used in their study were The bene®ts derived by E. ruidum from the use of this
presented pinned to the substrate and weight was not ®nely graded recruitment give an undeniable adaptive
considered as a parameter in this experiment since none value to the expression of such a strategy and probably
of the prey could be moved. By contrast, in the case of account both for the ecological success of this species in
E. ruidum, this parameter appears to be the key-factor the Neotropics and for its predatory impact on arthro-
allowing individual hunters to gauge the feasibility of pod populations (see Lachaud et al. 1996). Searching for
prey loading and, if recruitment is needed, to match the food is an energetically costly activity (Nielsen et al.
number of recruited workers. 1982; Lighton et al. 1987) and the cost of maintaining
In our experiments, chemical trail laying never oc- many workers out foraging is important. Nevertheless,
curred though it has been clearly demonstrated that such such a disadvantage may be outweighed by a suciently
trail laying from the Dufour's gland is involved in mass increased probability that food items are successfully
recruitment to rich or dicult food sources (Pratt 1989; retrieved (Mo€ett 1988). This is what occurs in E. rui-
Bestmann et al. 1995). The di€erences in the experimental dum, which is able to secure a high probability of pro-
procedure used by these authors may have a bearing both visioning success using di€erent ways. When solitary
on our failure to obtain mass recruitment with chemical strategies are involved, they can adopt an ecient
trail-laying (Lachaud 1985; Schatz et al. 1996; and this cleptobiotic behavior, either using the foraging trail laid
study) and on Pratt's failure to ®nd graded recruitment in by other species to take food items from them when they
E. ruidum. The same procedure as that used for P. clavata are returning to their nest (Perfecto and Vandermeer
(Breed et al. 1987) was employed by Pratt (1989) and 1991) or entering nearby colonies of the same species to
Bestmann et al. (1995) to trigger recruitment (i.e., large, steal their stores (Breed et al. 1990, 1992). When col-
unwieldy, freshly killed prey, pinned to the ground). lective strategies are involved (this study), their eciency
Consequently, the prey were ``in®nitely'' heavy to the is obvious since the majority of prey which trigger re-
scout ant which probably adopted a maximal response in cruitment are brought back to the nest (88 out of 94
its recruitment and so displayed the more ecient trail- prey). In collective strategies, the cooperative retrieving
laying behavior. In the same way, though two sizes of of the prey apparently does not imply a reduction in
protein bait (whole beetles, Pelidnota sp., versus beetle transport costs, which may even be increased by inter-
pronota only) were compared by Pratt in order to test the ference between ants and ineciency in group transport
ability of E. ruidum to match numbers of recruited (Sudd and Franks 1987). As reported by Pratt (1989),
workers to the sizes of baits, both prey were of similar when two or more E. ruidum workers cooperate in re-
unwieldiness and both probably triggered the same un- trieving a large prey item, they often worked at cross-
348

purposes and generally proceed very slowly. However, previous draft of the manuscript and to T. Williams for improving
Duncan and Crewe (1993) suggested that the extra en- the English text. This research was supported by grants of the
``Science de la Cognition'' (MESR) to B.S., the ``Cognisciences''
ergy needed for more foragers is probably compensated (CNRS) and ``PRESCOT'' programs to G.B. and J.P.L., the
by the energy obtained from the larger food, an hy- ``Conseil ReÂgional Midi-PyreÂneÂes'' to G.B. and the research pro-
pothesis con®rmed by our laboratory results. For short gram 0574P-N from CONACYT to J.P.L.
food±nest distances, the energetic bene®ts (expressed in
mean weight of prey retrieved per individual) derived by
E. ruidum from the use of recruitment systems are en-
hanced in comparison with the solitary hunting. Fur- References
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