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Sitemate Recognition: the Case of Anochetus Traegordhi (Hymenoptera; Formicidae) Preying on Nasutitermes (Isoptera: Termitidae) by B

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Sitemate Recognition: the Case of Anochetus Traegordhi (Hymenoptera; Formicidae) Preying on Nasutitermes (Isoptera: Termitidae) by B 569 Sitemate Recognition: the Case of Anochetus traegordhi (Hymenoptera; Formicidae) Preying on Nasutitermes (Isoptera: Termitidae) by B. Schatz', J. Orivel2, J.P. Lachaud", G. Beugnon' & A. Dejean4 ABSTRACT Workers of the ponerine ant Anochetus traegordhi are specialized in the capture of Nasutitermes sp. termites. Both species were found to live in the same logs fallen on the ground of the African tropical rain forest. A. traegordhi has a very marked preference for workers over termite soldiers. The purpose of the capture of soldiers, rather than true predation, was to allow the ants easier access to termite workers. During the predatory sequence, termite workers were approached from behind, then seized and stung on the gaster, while soldiers were attacked head on and stung on the thorax. When originating from a different nest-site log than their predator ant, termites were detected from a greater distance and even workers were attacked more cau- tiously. Only 33.3% of these termite workers were retrieved versus 75% of the attacked same-site termite workers. We have demonstrated that hunting workers can recognize the nature of the prey caste (workers versus termite soldiers) and the origin of the termite colony (i.e. sharing or not the log where the ants were nesting), supporting the hypothesis that hunting ants can learn the colony odor of their prey. This, in addition to the nest-site selection of A. traegordhi in logs occupied by Nasutitermes can be considered as a first step in termitolesty. Key words: Anochetus, Nasutitermes prey recognition, predatory behavior. INTRODUCTION During their 100 million years of coexistence, ants and termites have been engaged in a coevolutionary arms race, with ants acting as the aggressor and employing many predatory strategies while termites are the prey presenting several defensive reactions (1-11511dobler & Wil- 'LEPA, CNRS-UMR 5550, Universite Paul-Sabatier, 118 route de Narbonne, 31062 Toulouse cedex, France (e-mail: schatz©cict.fr)(correspondence author: B. Schatz) 2LEEC, CNRS-UMR 2214, Universite Paris-Nord, ay. J.B. Clement, 93430 Villetaneuse, France 3ECOSUR, Apdo Postal 36, 30700 Tapachula, Chiapas, Mexico 4LET, CNRS-UMR 5552, Universite Paul-Sabatier, 118 route de Narbonne, 31062 Toulouse cedex, France 570 Sociobiology Vol. 34, No. 3, 1999 Schatz, B. et al. - Prey Recognition in Anochetus 571 son1990). Even if a large percentage of ant species prey on termites Table 1: Coexistence of Nasutitermes termites colonies with Anochetus traegordhi and when given the opportunity, the Ponerinae subfamily seems to be one Odontomachus troglodytes colonies in logs. of the most specialized, from both a behavioral and morphological Sites studied standpoint, in their capture (Wheeler 1936; H011dobler & Wilson 1990). Ok Ants from three subfamilies (the ponerine genera Anochetus and Rotten logs with Ottotomo Bertoua Abong Matomb Nzi Total Odontomachus, the formicine genus Myrmoteras and the myrmicine tribe Dacetini with 24 genera) have each independently acquired a trap- Mbang Nasutitermes colonies 21 8 4 4 2 39 / jaw system that has emerged in response to the particular ecology of these ants (reviewed in Gronenberg & Ehmer 1996). Anochetus spp. are Anochetus traegordhi 17 6 3 4 1 31 79.5 encountered all around the world in tropical zones covering North, 12.8 Central and South America (Wheeler 1936; Mill 1982; Wilson 1987; Odontomachus troglodytes 2 1 1 0 1 5 Olson 1991; Jaffe & Lattke 1994; Longino & Hanson 1995; Dejean & Without ant colonies 2 1 0 0 0 3 7.7 Olmstead 1997), Australia (Wheeler 1936; Andersen & Clay 1996), Asia (Gronenberg & Ehmer 1996) and Africa (Levieux 1972; Villet etal. 1991), origin of the prey (same versus different from the ants). where they have been generally recorded as specialized termite preda- Comparisons were made using the Fisher's exact test (StatXact 2.05 tors. Some species have been reported to be specialized on a few termite software). species (Wheeler 1936; Mill 1982). RESULTS We study here the predatory behavior of the African A. traegordhi specialized in the capture of Nasutitermes (Dejean et al. 1999). We Nest site selection compared the predatory sequences of foraging workers confronted with Colonies of Nasutitermes termites were found in 39 logs, 92.3% of Nasutitermes workers and soldiers from colonies sharing or not the logs which were also inhabited by an Odontomachini subtribe ant colony where we found the ant colonies. OETECTION OF PREY) Q)PTEC7TION OF PREY) A. B. MATERIALS AND METHODS 73.2% ...11116. 26.8% 41111••• (LOCALIZATION) (LOCALIZATION) The ants and termites originated from the Democratic Republic of the 4111, 2.6% Congo (Kikwit and Kinshasa) and from Cameroon (Abong Mbang, ( APPROACH) 10.11% CAPPROACH) 1163.3% 10.1% Bertoua, Ottotomo, Matomb, Nzi). Numerous rotten logs lying on the (3.7 % , soil were examined to detect the simultaneous presence of colonies of 193% (ANTENNATION) (ANTENNATION) 12.2% 419.5% A. traegordhi and of Nasutitermes. 411111." 68.3% ( ATTACK) Observations on predatory behavior were carried out according to a 1,14 13% 6.8% 44.46. method developed during studies conducted on Dacetini (Dejean 1987; 26.3% 12.6% 11.4% (8ELFGROOMINC) CPREY MOVING) (FREY MOVING) see also Dejean & Everaerts 1997). A preliminary study allowed us to 4111- determine the complete behavioral sequence used by hunting ants (uFrit•,T) (`TRAc▪rio (LIFTING) (TRACTI4 .x,21.0%_/_ 3% 411. 8.8ZA4.6% when confronted with termites. Index cards were compiled with the 43.94% (STINGING) 34.2% QTINGING., respective postures of the predators and the prey noted for each phase hp 273% ► he 24.0% TRANSPORT" \ 40). (PREY MOVING) 33.0% CRANSPORI),), (FREY MOVING) of the sequence. We compared the predatory behavior of A. traegordhi 9.8% OF THE PREY)4 12.7% i. OF THE PREY 12.7% 4. when confronted with Nasutitermes workers and soldiers nesting in the ( %.(NEW STINGING) 53? 411k same log as the ant colony, or collected from a different log, using flow- r PREY ) CNTRY INTCh ( PREY (ENTRY INTO diagrams illustrating the capture of these four types of prey. Percent- \ABANDONE THE NEST) PREY ESCAPE) BAN DONE THE NEST (PREY ESCAPE) 59.5% 25.3% 15.2% ages (transition frequency between behavioral acts) were calculated 15.1% 65.9% 19.0% from the overall number of cases. We tested therefore the prey caste Fig. 1. Flow diagram of the A. traegordhi hunting workers when confronted with termite workers effect (workers versus termite soldiers) and the influence of the nest-site from the same log (A) (n = 205) or from a different nest site (B) (n = 79). 572 Sociobiology Vol. 34, No. 3, 1999 Schatz, B. et at — Prey Recognition in Anochetus 573 Percentage OETECTION OF PREY) B (pETECTION OF PREY) 80 80 A 45,733% 45, 87.5% (LOCALIZATION) (LOCALIZATION) 60 60 4111, 12.5% 45, APPROACH) APPROACH) ` 40 40 PASSIVE PASSIVE 45,73.5% 41'85.0% REY ESCAP (REY ESCAP) (ANTENNATION) (ANTENNATION) 20 20 IVO% 1.0% 6.0% ( ATTACK) C:-rrAci)11111683% 63% 0 PREY ESCAPE 10.0 0 PFTREERY DESCAPENSD 4.25.0% (WIER DEFENS (A 80 80 2.5%4 (LIFTINCD NI 60 (PREY MOVIN 60 '4,19.0V (iTINGINO 5.0% 40 40 6.0%i 1,16.5% CRANSPORTy-ir (---PREY ND TRANSPORT 20 20 O F THE PREY 1 0% OF THE PREY 3.5, . Ns ; •Ip 19.0% csigEW STINGING 9.4% (ENTRY ma\ PREY ) (ENTRY 1NT Head Thorax Gaster Head Thorax Gaster \,.. THE NEST,/ (PREY ESCAPE (tHANDONE THE NEST C PREY ESCAPE) 66.0% 19.0% 15.0% 75.6% 9.4% 15.0% 111 Approach Worker termites Soldier termites Fig. 2. Flow diagram of the A. traegordhi hunting workers when confronted with termite soldiers Attack from the same log (A) (n = 200) or from a different nest site (B) (n = 160). Fig. 3. Part of prey body used as a target during approach and seizure phases with termite workers living in the same log as their predator ants (n = 180) (A) and from a different nest site (n = 60) (Table 1). Four out of five logs (79.5%) were inhabited by A. traerghordhi (B), and with termite soldiers from the same log (n = 166) (C) and from a different nest site (n = and one out of eight logs (12.8%) by Odontomachus troglodytes. No case 130) (D). of coexistence of both ant species in the same log was recorded. worker. Whatever the type of prey, the hunting workers used the same Common traits of the predatory sequences for the four types of general sequence of predatory behavior. Nevertheless, the duration of prey (Fig. 1 & 2) each predatory phase and some qualitative patterns varied according During their search for prey, the A. traerghordhi workers moved to prey caste and nest site. slowly and sinuously, antennae spread apart and mandibles opened to Effect of prey caste 180°. Prey were detected from a distance of 5 mm in at least 73% of the The effect of prey caste (workers versus termite soldiers) was cases. This initiated the phase of localization followed by a very slow observed for each phase of the sequence of predation, except for the approach with the ant generally pointing its antennae towards the prey position of the antennae and the speed of mandible closure. Even if rather than keeping them bent. When prey were detected by contact, an generally both castes were approached very slowly, termite soldiers attack rapidly followed after a short antennation. The attack consisted triggered a slow approach more often than workers. They also triggered in the very rapid closure of the mandibles (the trap jaw system). The shorter antennation only when they originated from the same logs as prey was sometimes projected several centimeters away. In some cases, the hunting workers.
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