Evaluation of the Diet of Odontomachus spp. on the Island of Dominica

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1 Evaluation of the Diet of Odontomachus spp. on the Island of Dominica Samuel Ryan Shook Texas A&M University June 2017

2 1 Abstract: In 2015, a second species of ant in the genus Odontomachus, O. ruginodis, was discovered to be living on the island of Dominica alongside the originally known species O. bauri. Both are highly predatory species known for developing close associations with and preying on termite nests, although their dietary preferences outside of that are relatively unknown. The purpose of this study was to catalogue the prey intake of O. bauri and O. ruginodis nests under different conditions, so that that data can be used to examine aspects of the life history of each species, and compare the two. The prey taken by foragers was recorded by observation of foragers returning to the nest, either identifying it as the worker carried it in or retrieving it from the worker for later identification. It was found that colonies of either species that were associated with termite nests took termites as prey to the exclusion of all other insects, but species living away from them acted as generalist predators and scavengers focusing on soft-bodied arthropods. Introduction: Odontomachus is a pantropical genus of ants in the subfamily Ponerinae, commonly called Trap-jaw ants. They are large ants, with their most notable feature being their large mandibles, which can be opened wide and then locked into place (Fowler 1980). Upon encountering a prey item, the mandibles are unhooked, and swing forward at high speeds to stun or latch onto prey, the eponymous trap jaw (Dejean and Bashingwa 1985). Until recently, it was thought that the only species in the genus on the island of Dominica was Odontomachus bauri. However, in 2015 Andrew Graf discovered a nest of Odontomachus ruginodis, a similar species to O. bauri (Graf 2015). Both are ground-dwelling species, which are often found in close association with termite nests (Graf 2015). While a number of studies have been done on the association of trap-jaw ants in the genus Odontomachus and termites, including their habit of preying on them, little has been done with their predation of other groups of insects. They are known to take prey other than termites, although the diversity of prey has not been extensively investigated, and were noted in one study as regularly scavenging, although this was observed only rarely by other studies (Holldobler and Ehmer 1995, Camargo and Oliveira 2012). This leaves room for both confirmation of what has already been observed, and further exploration of facets of the lives of the ants of both species that have not been studied in detail. The purpose of this study is to examine the diet breadth of nests of Odontomachus bauri and Odontomachus ruginodis on the

3 2 island of Dominica, cataloguing the types of insect prey being brought in by foragers, the number of each type of prey, and if possible whether the prey was captured live or scavenged. The differences in prey between nests can then be compared, examining differences in preference between O. bauri and O. ruginodis, between nests in different habitats, and between nests that are associated with termites or free living. Materials and Methods: Data collection was performed from May 27 th to June 8 th, 2017, on the island of Dominica in the West Indies. For each nest, the GPS coordinates (taken either with a Garmin 78 GPS or the app MotionX-GPS), date of collection, species, and location of the nest in the environment was noted. Data were collected by observing a nest until a forager returned with prey. Once a laden forager was spotted, the prey was either removed from the forager s mandibles with forceps or the prey and forager were collected with an aspirator. If the prey item couldn t be recovered, but was definitively identified while the forager was carrying it, then it would also be recorded. Six nests were collected from, as follows: Nest 1: Roseau Botanical Gardens, under a termitarium, O. bauri N W Nest 2: Roseau Botanical Gardens, under the roots of a tree, O. ruginodis N W Nest 3: Springfield Research Station, under the roots of a tree, O. bauri N W Nest 4: Cabrits National Park, under a termitarium, O. ruginodis N W Nest 5: Kalinago Territory, under the roots of a tree, O. bauri N W Nest 6: Springfield Research Station, under a tree stump, O. bauri N W Specimens were identified as either Odontomachus bauri or O. ruginodis by comparison to images of species in the genus Odontomachus found on Antweb (Antweb 2017).

4 Table 1. Number of prey of each type taken at each nest, and total number of each type overall. 3 Termite Alates Termite Workers Caterpillars Crickets Ants Isopod Hemipteran Millipede Pupa Spider Earwig Diptera Nest 1 3 Nest Nest Nest Nest Nest Total Results: Fig 1. Proportion of each prey type taken from all nests. Termites make up a plurality of all prey items taken at 37% of the total, with the nests associated with termitaria (Nests 1 and 4) being observed only taking termite prey. Nests not directly associated with a termite colony were never observed with termite workers, only collecting termite alates after a nuptial flight (Table 1). The next most common prey

5 4 item was caterpillars at 19%, followed by ants at 12% and spiders at 10% (Figure 1). Although the state of the majority of the prey at time of capture was not observed, in two instances a forager was seen taking food that had clearly been dead for some time, these being one half of a millipede at Nest 3 and an earwig and desiccated isopod at Nest 6. With the exception of the ants and the non-scavenged isopod, all mobile prey collected was soft-bodied, with the crickets and hemipterans all being immatures. Of the six ants collected, only two were workers, one from Nest 2 and one from Nest 6, with the other four being alates, three males and one female. Discussion: The preponderance of termite prey in the diet, particularly for those associated with termite nests is unsurprising, as there have been a number of studies that have observed this as well. What is more interesting is that those nests that were not built next to a termitarium were not observed with any termite workers as prey, only eating termites during mass flights of alates. This suggests that either the nests away from termitaria were taking termites from termite nests further away and just happened to not do so while being observed, or that these two Odontomachus species prefer termites to such a degree that when they are available the colony will build nests near termitaria when possible. One potential avenue of future research would be to look at the distribution of Odontomachus nests in areas with high and low densities of termitaria. The strong preference for soft-bodied prey was noted in another study (Holldobler and Ehmer 1995), but with a focus on preying on termites specifically rather than soft-bodied prey in general. However, personal observations of the Odontomachus foragers may provide an explanation. On discovering a prey item the foragers were noted as striking at the prey several times, retreating slightly with each one, until it would keep a hold on the prey after a strike and use its stinger, which matches previous observations of Odontomachus bauri hunting strategies (Holldobler and Ehmer 1995). When foragers were seen attacking prey with harder exoskeletons, it would often take a larger number of strikes before the forager would sting, with the prey often fleeing during the retreat between strikes. In contrast, soft bodied prey was often stung either on the first or within the first few strikes, leaving less opportunity for escape. If the forager retreats after each strike because it failed to get a firm grip on the prey and needs to reopen its mandibles for another strike, then it s possible that foragers can simply grip soft-bodied prey more easily and so find them easier to capture. The next most common prey item after termites were caterpillars,

6 5 making up a larger portion of the prey captured at nests 2, 3, and 5, all of which were under the roots of living trees. The location of these nests, coupled with the lack of caterpillars as prey at Nest 6, which was not underneath the branches of any nearby trees, makes it plausible that small caterpillars that had fallen from the tree above the nest were simply the most commonly available prey item. Other Odontomachus species have been shown to be generalist predators in the past, and so a diet consisting of whatever suitable prey happens to be nearby would make sense (Camargo and Oliveira 2012). Despite not being soft bodied, other ants and particularly ant alates were a common prey item, both in this study and in previous ones (Holldobler and Ehmer 1995, Camargo and Oliveira 2012). One potential explanation for this is that the abundance of prey offsets its lower quality. Ant alates, like termite alates, swarm in high numbers, and this overabundance of them leads to foragers encountering them so often that they will make up a significant portion of the diet even if many of them escape capture. In addition, male alates often die shortly after mating, and it is much easier to take those carcasses over even easy living prey (Shik and Kaspari 2009). Scavenging hard-bodied insects is something that at least Odontomachus bauri has been demonstrated as performing, both in this study and in at least one previous one, though that previous study was not specific about the relative proportion of captured vs scavenged food beyond that retrieval of dead items was repeatedly witnessed (Holldobler and Ehmer 1995). The presence of O. ruginodis on Dominica was only discovered in 2015 by Andrew Graf, and so a comparison of their diet preferences on the island has not been performed. According to the data found here, there does not appear to be a significant difference between the diets of O. bauri and O. ruginodis. Both are found associated with termites, eating largely or exclusively termites when associated with them, and otherwise have a preference for soft-bodied arthropods. There was no existing literature on the diet preferences of O. ruginodis like there was for O. bauri, which means there was no baseline of comparison for the accuracy of the data found. One major issue with the comparison between these two species, and with the study as a whole, is that time constraints limited the time spent observing nests in places other than Springfield research station. Since both nests observed on the station were O. bauri colonies, this means that there could be significant differences between O. bauri and O. ruginodis that weren t apparent in the limited observation window. In addition, while both species were found at the Roseau Botanical

7 6 Gardens, the O. bauri found there were associated with a termitarium, which has been shown to significantly alter their foraging. This means that all equitable comparisons between O. bauri and O. ruginodis are being made between nests in often significantly different habitats. Locating a colonies of two different species that are in close proximity and studying their dietary preferences without habitat as a factor could be an interesting avenue of future research. More than anything else, the biggest limitation of this study was available time. Only two of the nests, the ones at Springfield, could be observed over multiple days, with the others observed only once and often only briefly. In addition, all of these observations were made during the late morning and early to mid afternoon, which could be problematic if one or both species is primarily nocturnal or crepuscular. Previous studies have not established any such daily pattern in O. bauri, but other species in the same genus have been shown to prefer to forage at night (Holldobler and Ehmer 1995, Camargo and Oliveira 2012). In order to confirm the accuracy of this study, a future study could focus on repeatedly visiting a smaller number of colonies of each species under a wider variety of conditions. These are highly predatory ants, and so predation is one of the primary ways in which they interact with other insects, making understanding their habits as predators important to fully understanding their life history. By gathering information about the ants' feeding habits, it is hoped that this can act as a foundation on which a complete picture of the place of these ants in the insect community of the island can be created. Acknowledgements: I would like to thank Dr. Adrienne Brundage, Dr. Thomas Lacher, and Dr. Jim Woolley for their advice and assistance in developing and performing this experiment, and for allowing me the opportunity to come to Dominica in order to carry it out. Works Cited: Dejean A. & Bashingwa E La Predation Chez Odontomachus Trodlodytes Santschi (Formicidae-Ponerinae). E.P. Ins. Soc 32: 23. doi: /bf Andrew Graf, A survey of Odontomachus sp and General Observations of Nesting Preferences on the Island Nation of Dominica in the West Indies, 2015,

8 7 Andrew_2015.pdf Ehmer B. and Holldobler B Foraging Behavior of Odontomachus bauri on Barro Colorado Island, Panama, Psyche: Volume 102, Issue 3-4, Pages Camargo R.X. and Oliveira P.S Natural history of the Neotropical arboreal ant, Odontomachus hastatus: Nest sites, foraging schedule, and diet. Journal of Insect Science: Volume 12, Issue 48. Fowler H.G Populations, prey capture and sharing, and foraging of the Paraguayan Ponerine Odontomachus chelifer Latreille. Journal of Natural History, 14:1, 79-84, DOI: / Shik, J.Z. & Kaspari, M Lifespan in male ants linked to mating syndrome. Insect. Soc. 56: 131. doi: /s AntWeb. Available from: t=allantwebants. Accessed 10 June AntWeb. Available from: oject=allantwebants. Accessed 10 June 2017.

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