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2012 Observations on the Specialized Predatory Behavior of the Pitchfork-Mandibled Ponerine Ant Thaumatomyrmex Paludis

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2012 Observations on the Specialized Predatory Behavior of the Pitchfork-Mandibled Ponerine Ant Thaumatomyrmex Paludis US ISSN 0006-9698 CAMBRIDGE,MASS.2012NUMBER 533 OBSERVATIONS ON THE SPECIALIZED PREDATORY BEHAVIOR OF THE PITCHFORK-MANDIBLED PONERINE ANT THAUMATOMYRMEX PALUDIS (HYMENOPTERA: FORMICIDAE) {, CHRISTIAN RABELING 1,MANFRED VERHAAGH2, AND MARCOS V. B. GARCIA3 ABSTRACT. In this natural history note we describe and illustrate the specialized predatory behavior of the ponerine ant Thaumatomyrmex paludis from the Brazilian Amazon. This study of T. paludis implies that specialized predation on polyxenid millipedes is widespread in the genus Thaumatomyrmex. The observation that one T. paludis forager only partly depilated its prey before it started feeding suggests that complete depilation of polyxenids, as reported from T. contumax, could be most important when prey items are fed to the larvae, presumably to protect the brood from irritating bristles. Our observations provide an additional example of the sympatric distribution of two Thaumatomyrmex species, further supporting the hypothesis that the genus consists of multiple, morphologically similar species. RESUMO. Neste trabalho descrevemos e ilustramos o comportamento predato´rio especializado da formiga do grupo Ponerinae Thaumatomyrmex paludis da Amazoˆnia brasileira. Este estudo indica que o comportamento predato´rio especializado em diplo´podes da ordem Polyxenida e´ comum no geˆnero Thaumatomyrmex. Ale´m disso, nossas observac¸o˜es indicam que as formigas forrageiras depilam a presa apenas parcialmente logo apo´s a captura para consumo imediato. Depilac¸a˜o completa da presa, como descrito em T. contumax, parece ser mais importante quando a presa e´ usada para alimentar as larvas, sendo provavelmente praticada para a protec¸a˜o da cria contra cerdas irritantes. Este estudo ainda demonstra que uma outra espe´cie do geˆnero ocorre em simpatria com T. paludis. Estas observac¸o˜es corroboram a hipo´tese de que este geˆnero e´ composto por mu´ltiplas espe´cies morfologicamente similares. KEY WORDS: predation; ants; sympatry; mandible morphology; Isopoda; Polyxenidae { Corresponding author email address: crabeling@gmail. com INTRODUCTION 1 Museum of Comparative Zoology Labs, Harvard University, 26 Oxford Street, Cambridge, Massaschu- The mandibles of an ant are functionally setts 02138, U.S.A. analogous to the hands of a human. They are 2 Staatliches Museum fu¨r Naturkunde Karlsruhe, Erb- prinzenstr. 13, 76133 Karlsruhe, Germany. used to fulfill a variety of tasks such as nest 3 Embrapa Amazoˆnia Ocidental, Rodovia AM-10, Km building, defense against enemies, and ma- 29, Caixa Postal 319, Manaus, AM 69010-970, Brazil. nipulating brood (Ho¨lldobler and Wilson, E The President and Fellows of Harvard College 2012. Breviora brvo-533-00-01.3d 19/11/12 16:24:53 1 Cust # MCZ3 2 BREVIORA No. 533 1990). Most important, the mandibles are and chemically well-protected millipede spe- precision instruments for catching and han- cies, polyxenids are soft-bodied and rely on dling prey. Ants have evolved diverse preda- hooked bristles for defense (Eisner et al., tory behaviors, accompanied by an astounding 1978). When attacked, they wipe hairs from a variety of morphological specializations, in- pair of bristle tufts at the caudal end of their cluding the sickle-shaped mandibles of isopod- bodies against the aggressor and when the hunting Leptogenys (Brown, 2000), the filigree attacker attempts to clean itself, the barbed forceps-like mandibles of Martialis heureka hairs lock into the predator’s setae, interlink (Rabeling et al., 2008), and the trap jaws of with one another, resulting in the predator’s dacetine springtail hunters and of Odontoma- complete immobilization (Eisner et al., 1996). chus ants (Bolton, 2000; Patek et al., 2006). So far, only Neotropical Thaumatomyrmex The perhaps most astounding case of man- ants and the Indonesian ant species Probolo- dibular morphological specialization in ants is myrmex dammermani are known to be spe- the bizarre, pitchfork-shaped mandible of cialized predators of polyxenid millipedes the Neotropical ponerine ants in the genus (Branda˜o et al., 1991; Eisner et al., 1996; Thaumatomyrmex. The genus name, which Ito, 1998). translates approximately to ‘‘wondrous’’ or Our serendipitous finding of two rarely ‘‘marvelous’’ ant, emphasizes the outlandish collected Thaumatomyrmex species during appearance of these remarkable insects (Mayr, field studies in the Brazilian Amazon moti- 1887). vated this short natural history note. It Since the discovery of the first Thaumato- provides information on the predatory be- myrmex species no doubt existed that these havior of the Amazonian T. paludis and an ants must be specialized predators. Specula- additional example of sympatrically occur- tions on the most likely prey organisms ranged ring Thaumatomyrmex species. from snails to termites to the brood of other ant species (Creighton, 1928; Weber, 1942; Kempf, 1975). However, no actual predatory MATERIALS AND METHODS behavior was observed and therefore the In the late afternoon of 29 May 2003, a nature of the prey and the details about the single stray Thaumatomyrmex worker was function of Thaumatomyrmex’s mandibles collected at the experimental agricultural remained a mystery for over a century. Diniz field site of the Embrapa, Empresa Brasileira and Branda˜o (1989) finally solved the puzzle, de Pesquisa Agropecua´ria, in Manaus, Ama- reporting that Thaumatomyrmex ants are zonas, Brazil, located at kilometer 28 of specialized predators of bristly millipedes Highway AM 010 (26539S, 596599W; 40–50 m belonging to the family Polyxenidae (Diplo- poda: Polixenida: Polyxenidae; sensu Conde´ above sea level). The worker was presumably and Nguyen Duy-Jacquemin, 2008). These foraging in the thick leaf litter layer of an first observations were complemented by agroforest, characterized by a diverse mix of careful laboratory experiments on T. contumax fruit and nut trees. Thaumatomyrmex colo- and field observations on T. atrox from the nies are of minute size, sometimes containing Brazilian state of Bahia (Branda˜o et al., 1991). fewer than five workers (Branda˜o et al., Additional feeding experiments conducted for 1991; Delabie et al., 2000; Jahyny et al., T. contumax confirmed these results (Delabie 2002). Our efforts to locate the colony failed, et al., 2000; Jahyny et al., 2008). and we kept the single forager alive in a Petri Polyxenid millipedes represent an unusual dish to which we added leaf litter and a moist prey, because in contrast to heavily sclerotized cotton swab to provide humidity. No food Breviora brvo-533-00-01.3d 19/11/12 16:24:55 2 Cust # MCZ3 2012 PREDATORY BEHAVIOR OF THAUMATOMYRMEX PALUDIS 3 was added to the formicarium until the start Thaumatomyrmex remains utterly confused of the feeding experiments on 31 May 2003. andisindireneedofacomprehensive For the feeding trials, the ant was transferred revision. Measurements and indices for the into a watchmaker’s glass dish and different second unidentifiable species, hereafter re- arthropods such as Drosophila flies, iso- ferred to as T. sp. Manaus 2, are: WL pods, termites, and polyxenid millipedes (0.83), HL (0.55), HW (0.61), ML (0.69), SL were added to the arena. Photographs of (0.47), CI (111), SI (86), MI (125). The the foraging worker were taken with a Nikon specimens were deposited at the Museu de Coolpix 995, which was handheld to a Wild Zoologia da Universidade de Sa˜o Paulo in M-5A stereomicroscope. Brazil and at the Staatliches Museum fu¨r In June 2003, we collected a second T. paludis Naturkunde in Karlsruhe, Germany. Indi- worker at the same agroforest site, carrying a viduals were imaged at the Museum of polyxenid millipede between its mandibles, Comparative Zoology using a Leica presumably returning to the nest. In addition, DFC420 digital camera mounted on a Leica we collected a single worker of a second MZ16 dissecting scope. Composite images Thaumatomyrmex species in an adjacent patch were assembled using the Leica Application of primary terra firme rainforeston16April Suite (v. 3.7.0) and Helicon Focus (v. 5.2.9) 2003. We were not able to find the colonies of software packages. these ants either, and we did not conduct laboratory experiments with these individuals. RESULTS To identify the individuals to species level, we used taxonomic keys published by Kempf To study and document the predatory (1975) and Jahyny et al. (2008). Morpho- behavior of T. paludis, we first placed the metric measurements were taken as defined single worker into the foraging arena and in Kempf (1975) and are given below. The subjected the individual to a cafeteria-style two workers collected at the agroforest site feeding experiment. First, a termite worker were identified as T. paludis Weber, 1942 (as was added. The forager antennated the resurrected from synonymy by Jahyny et al., potential prey briefly, turned away, and 2008; see also Longino, 1988) on the basis of continued foraging in the arena. Subsequent- the absence of the metanotal suture and ly, a Drosophila fly and an isopod were the relatively short mandibles (Fig. 1). The presented to the Thaumatomyrmex worker, morphometric measurements (given in mm) soliciting the same response: brief antennat- and indices (used as defined in Rabeling ing followed by disregard. Only the release of et al., 2007) for the two T. paludis individuals a polyxenid millipede into the foraging arena are: WL (0.97, 1.02), HL (0.65, 0.65), HW immediately solicited predatory behavior (0.63, 0.61), ML (0.61, 0.59), SL (0.57, 0.55), from the T. paludis worker, which followed CI (97, 94), SI (88, 85), MI (94, 91). The third the characteristic pattern described for T. Thaumatomyrmex individual, collected in a contumax (Branda˜o et al., 1991). The milli- primary rain forest, could not be identified pede was first antennated by the forager, with certainty, but might be related to T. then briefly secured between the mandibles, soesilae Makhan, 2007 (Fig. 1). However, stung, and released immediately afterward. the taxonomic identity of T. soesilae is The Thaumatomyrmex worker retreated and uncertain, given the inadequacy of the the millipede did not attempt to fling its original description and the unavailability barbed hairs against the attacker.
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