2013 SOUTHEASTERN NATURALIST 12(2):367–378 Ant Species in the Diet of a Florida Population of Eastern Narrow-Mouthed Toads, Gastrophryne carolinensis Mark Deyrup1,*, Leif Deyrup2, and James Carrel3 Abstract - Gastrophryne carolinensis (Eastern Narrow-mouthed Toad) is known to be an ant specialist, but prey identification has rarely progressed beyond family level. There are no prey records from Florida scrub, a rare upland habitat type. This study identifies species of 4859 individual ants retrieved from stomachs of 146 G. carolinensis collected in Florida scrub. All toads had consumed ants; ants comprised about 95% of all food items. Forty-three species of ants were recorded. About 77% were various species of Pheidole or Nylanderia. The ants consumed were mostly small (4 mm or less in length) and nocturnally active. Species that were eaten belong to ant genera known to contain venoms, chemical repellents, or other organic substances in exocrine glands. This finding suggests the possibility that Narrow-mouthed Toads have opportunities to sequester exocrine secretions of ants, in the manner of some other anurans. The diversity of ant species consumed by G. carolinensis suggests that this species might be able to subsist on disturbed-site ants, including exotic species such as Solenopsis invicta. Introduction Gastrophryne carolinensis (Holbrook) (Eastern Narrow-mouthed Toad) is a common amphibian found throughout Florida, with a range extending north into Maryland and west into Texas and Kansas (Nelson 1972). It may be the most numerous amphibian species at some southeastern sites (Dodd et al. 2007, Farrell et al. 2011, Tuberville et al. 2005). It is a small species, with an adult snout– vent length (SVL) of 23–27 mm (Nelson 1972). The Narrow-mouthed Toad is completely nocturnal, spending the day in its burrow or under debris (Anderson 1954, Dickerson 1906). The diet of North American Gastrophryne species has long been known to consist primarily of ants (Anderson 1954, Brown 1974, Carpenter 1954, Dickerson 1906, Fitch 1956, Garton and Mushinsky 1979, Nelson 1972, Tanner 1950). Fitch (1956) even called G. olivacea (Hallowell) (Western Narrow-mouthed Toad) “the Ant-eating Frog”, although this common name did not gain wide currency. Nelson (1972) suggests that Gastrophryne species show several adaptations for myrmecophagy, including a fold of skin that can be moved forward to cover and protect the eyes, a small mouth, especially compared with hylid frogs of the same size, and a tough skin that produces a sticky defensive secretion. Our study aimed to go beyond reaffirming that ants are the primary prey of G. carolinensis. Identification of ant species in stomach contents should permit 1Archbold Biological Station, 123 Main Drive, Venus FL 33960. 2Department of Biology, University of the Cumberlands, Williamsburg, KY 40769. 3Division of Biological Sciences, University of Missouri-Columbia, MO 65211. *Corresponding author - mdeyrup@ archbold-station.org. 368 Southeastern Naturalist Vol. 12, No. 2 correlations between Narrow-mouth Toad feeding behavior and size, daily activity patterns, defenses, and other specific features of prey species. Since the ant fauna of the study site is well known (Deyrup and Trager 1986, Wiescher 2010), it should also be possible to list some available species that are not included in the diet of G. carolinensis. Finally, this study should provide a dietary baseline, as the study site is a natural habitat with few non-native ants; such sites are becoming less frequent in peninsular Florida, a region heavily invaded by exotic ants (Deyrup et al. 2000). Although the primary purpose of this study is a detailed account of the diet of G. carolinensis, it also adds unusually detailed information on ants as dietary items. This latter subject is generally neglected by myrmecologists, who tend to be enthralled by the accomplishments of ants and not disposed to view ants as victims. A recent, comprehensive book on the ecology of ants (Lach et al. 2010), for example, does not examine the ecological importance of ants as the dietary mainstay of a wide variety of vertebrates and arthropods. Study Area The primary study site was the Archbold Biological Station (ABS) (27°11'N, 81°21'W), a 2001.6-ha private research station 12 km south of the town of Lake Placid in Highlands County, FL. The habitat where toads were collected is native Florida scrub, a shrub habitat that occurs on deep, well-drained sandy uplands. This habitat is kept open by periodic fires; for a general description of Florida scrub habitat and its ecology, see Myers (1990). Trapping of G. carolinensis was in two Florida scrub plant associations. One of these was scrub rosemary “balds”, a habitat dominated by Ceratiola ericoides Michaux (Sandhill Rosemary) shrubs, with patches of open sand between the shrubs. The other was “scrubby flatwoods”, dominated by scrub oaks, especially Quercus geminata Small (Sand Live Oak), Q. chapmanii Sargent (Chapman’s Oak), Q. inopina Ashe (Sandhill Oak), Serenoa repens (Bartram) Small (Saw Palmetto), Sabal etonia Swingle ex Nash (Scrub Palmetto), and ericaceous shrubs, such as Lyonia species. These uplands are interspersed with areas of flatwoods and seasonal ponds that have a high watertable in the rainy season. For a detailed description of habitats on the ABS, including plant associations, soil types, nutrients, soil moisture, and prevalent soil and air temperatures, see Abrahamson et al. (1984). Methods Gastrophryne carolinensis, 146 specimens, were captured incidentally in a study of burrowing wolf spiders, whose population dynamics are the subject of long-term studies by one of the authors (J. Carrel). Toads were collected in 15-cm-diameter, plastic, water-filled dessert bowls that were buried to their rims in the surrounding sand. The protruding rims appear to exclude most ground lizards, but G. carolinensis occasionally fell in and drowned. This species is able to float in water in breeding ponds (Dickerson 1906), and we assume that toads were affected by the presence of a small amount of dish-washing detergent, used 2013 M. Deyrup, L. Deyrup, and J. Carrel 369 to keep spiders from using surface tension to escape from the bowls. Most collections were made in March–November of 2007–2008, with the largest numbers of toads found in August–September. Specimens from an array of traps were combined over a 5-day interval and stored in 70% isopropyl alcohol. For example, a long series is labeled “5–9 August 2008, Ceratiola bald”. It is not possible at this point to assign exact sites and dates to specimens. Each toad was measured (SVL) and given a number prior to removal of stomach contents. Arthropods were surveyed from stomachs only, as intestinal contents were often too degraded to allow identification of prey species. Results Our examination of 146 G. carolinensis stomach contents showed that ants are the principle prey of G. carolinensis on the ABS (Appendix 1). All individuals had ants in their stomachs, and numbers of ants per stomach ranged from 1–313, with a mean of 33.3 and a median of 27.5. Although non-ant prey items occurred in most stomachs, the 4869 individual ants comprised about 95% of all prey items. Ants clearly made up most of the weight and volume of prey items, since 160 of the 266 additional food items were mites that are less than half the size of the smallest ant. The remaining non-ant insects were small Coleoptera, Hymenoptera, Heteroptera and Diptera, the largest of which (the elaterid Conoderus bellus) is only about 3.5 mm in length. Our study supports earlier findings of an ant-based diet for Gastrophryne species (Anderson 1954, Fitch 1956, Tanner 1950). The primary contribution of our study to the natural history of G. carolinensis is the species-level identification of prey, especially the 43 species of ants (Appendix 1). Discussion There is no reason to suspect that G. carolinensis has nutritional requirements that can only be met by particular species or genera of ants. Most of the species that we recorded are absent from the northern range of G. carolinensis, and are also absent from some habitats, such as cypress swamps (M. Deyrup, unpubl. data), where G. carolinensis may occur (Anderson 1954). At the generic level, all the ants we recorded are widespread in the southeastern and mid-Atlantic states, with the exceptions of Odontomachus and Hypoponera. It is possible that some of these genera have a special nutritional significance for G. carolinensis, but there is no evidence of this, and the diversity of ant genera represented in the stomach contents suggests a lack of selectivity. There are, however, large differences in the numbers of ants representing various genera in this sample of stomach contents. About 77% of all ants were in the unrelated genera Pheidole (consumed by 89% of the Gastrophryne) or Nylanderia (consumed by 76% of the Gastrophryne). The simplest working hypothesis to explain this is that members of these two genera were most accessible and vulnerable to pred ation. Gastrophryne carolinensis is known to be exclusively nocturnal (Anderson 1954, Dickerson 1906). As expected, ants found in the ABS sample of stomach 370 Southeastern Naturalist Vol. 12, No. 2 contents (Appendix 1) are generally species that are nocturnal or crepuscular during the summer at the ABS. Among the prey were two species of ants that are almost completely diurnal, Forelius pruinosus and Monomorium viride (M. Deyrup, unpubl. data). These, however, were rare in stomach contents— the former species consumed by one G. carolinensis, the latter by three. If G. carolinensis take up residence in or adjacent to ant nests, as reported by several observers elsewhere (Anderson 1954, Fitch 1956, Tanner 1950, Wood 1948), one would expect some toads would be associated with diurnal ants. At the ABS, there is no evidence that G. carolinensis moves into ant nests. Not only are there no records of large numbers of diurnal ants in stomach contents, but there are very few of the teneral (partially pigmented) adult ants that are usually found in ant nests. Most stomachs contained relatively small numbers of several species of ants, as if toads were consuming stray ants and small groups of ants foraging on the surface or in leaf litter. In the habitats where toads were sampled, most terrestrial ants nest deep in the soil or have diffuse nests in sand or leaf litter; the option of living in or adjacent to an ant nest may not be available. At the ABS, it appears that G. carolinensis can find sufficient prey without moving into an ant nest. Gastrophryne carolinensis appears to avoid ants in the genera Odontomachus (three specimens eaten) and Camponotus (two specimens eaten). These ants are largely nocturnal (Wiescher 2010). Members of both genera are common in the habitats where trapping occurred (Wiescher 2010). It is possible that these ants are too large to be readily acceptable: O. relictus is about 7 mm in length, while the Camponotus species range about 6–9 mm. The largest ants occurring in large numbers in stomach contents were species of Aphaenogaster, which are about 4–6 mm in length. Odontomachus workers have a powerful sting, while Camponotus can spray defensive chemicals. Dorymyrmex is another genus that might be underrepresented as prey. Nineteen G. carolinensis consumed a total of 50 Dorymyrmex (Appendix 1), but these ants are abundant in the study area and sometimes active at night during summer (Trager 1988, Wiescher 2010). Ants eaten by G. carolinensis almost certainly produce diverse medleys of exocrine secretions. Comparative studies over the past 40 years reveal that ants exude more than 140 organic substances from as many as nine types of exocrine glands (Morgan 2008). Venom glands of ants, the type of glands that has received the most intensive analysis, are remarkably versatile “factories” for natural products (Fig. 1; Blum 1992, Schmidt 1986). Ants with functional stings include species of Hypoponera, Odontomachus, Neivamyrmex, Monomorium, Pyramica, Solenopsis, and Strumigenys. Ants whose venoms have been characterized include species of Brachymyrmex, Nylanderia (Saporito et al. 2004, Smith and Jones 2004), Camponotus (Eisner et al. 1993), Dorymyrmex (Blum and Warter 1966, McGurk et al. 1968), Forelius (McGurk et al. 1968), Aphaenogaster (Wheeler et al. 1981), Odontomachus (Wheeler and Blum 1973), Solenopsis (MacConnell et al. 1970), Solenopsis (Diplorhoptrum) (Blum 1981), Trachymyrmex (Crewe and Blum 1972), and Cyphomyrmex (Crewe and Blum 1972). 2013 M. Deyrup, L. Deyrup, and J. Carrel 371 The prosperity of G. carolinensis through its native range will depend largely on adequate larval habitat (Dixon et al. 2011) and on the ability to persist in disturbed habitats dominated by exotic ants. Exotic ants in our sample are represented by six species, of which only two, Pheidole moerens and Solenopsis invicta, were consumed by more than three individual toads. This probably reflects the relative scarcity of exotic ants in undisturbed Florida scrub habitat (King and Porter 2007, Wiescher 2010). At the level of myrmecological surveys or the level of populations of foraging G. carolinensis, natural habitats in the southeastern coastal plain often persist only in a matrix of disturbed habitats such as improved pastures, fields, urban and suburban areas, and road edges (King and Porter 2007). Exotic ants in the Southeast are largely confined to disturbed Figure 1. Ants, arranged by subspecies and characteristic venoms, in the diet of Eastern Narrow-mouthed Toads in Florida scrub habitat. Diagram of families adapted from Tsutsui et al. (2008). 372 Southeastern Naturalist Vol. 12, No. 2 areas, where they may be a major part of the ant fauna (Deyrup et al. 2000, King and Porter 2007, King and Tschinkel 2006, Tschinkel 2006). In heavily disturbed sites, such as improved pastures, non-native ants comprise up to 90% of the total number of ants (King and Tschinkel 2006). The speciose ant diet that we documented at the ABS suggests that G. carolinensis is a generalist consumer of small ants and could subsist on a diet of small exotic ants. Almost all small exotic ants in Florida belong to genera represented in our sample of stomac h contents. The abundant Solenopsis invicta Buren (Red Imported Fire Ant) may present a special challenge or opportunity for G. carolinensis. Under certain conditions S. invicta can mass attack and kill small amphibians that have poor mobility (Todd et al. 2008, Tschinkel 2006). This ant has been frequently invoked to explain the decline of some vertebrates, including amphibians, but credible evidence of population-level effects is scarce (Tschinkel 2006). Most suggestions of such population declines are based on anecdotes, loose correlations, or situations where a variety of environmental influences are at play, only one of which is high populations of S. invicta (Tschinkel 2006). It is not unreasonable to suggest that the invasion of S. invicta may have benefitted G. carolinensis. The major benefit would be the introduction of an enormous dietary resource, consisting of ants of suitable size, usually found in shallow nests that erupt with potential victims when disturbed. This idea is supported by the presence of S. invicta in our samples and by reports of G. carolinensis in nest mounds of S. invicta (Garton and Mushinsky 1979). On the other hand, it is possible that an almost exclusive diet of S. invicta is unsuitable for G. carolinensis, or there might be a period of vulnerability to S. invicta in the life cycle of G. carolinensis, such as just after metamorphosis. There are many sites in Florida appropriate for examining the relationship between S. invicta and G. carolinensis. It might be difficult to adequately demonstrate population-level relationships between the two species, but it should be relatively easy to show whether G. carolinensis experiences S. invicta as a resource, or as a hazard, or perhaps both. The 266 prey arthropods other than ants comprise 5.2% of the individual arthropods consumed. The majority of non-ant arthropods are hard-bodied mites such as Oribatidae. These prey are under 1 mm in length and generally heavily armored. Oribatid mites appear to be a dietary source of protective alkaloids for dendrobatid “poison frogs” (Saporito et al. 2007). Some dendrobatid frogs have been described as “ant-mite specialists” (Saporito et al. 2007). Other non-ant arthropod prey items include a wide spectrum of small ground-dwelling species, none of which was eaten by more than a few toads. The diversity of non-ant prey appears to show a great flexibility in consumption of opportunistic prey. Gastrophryne carolinensis is an ant specialist and probably has specific ant-hunting behavior, but its wide range of alternative prey suggests that G. carolinensis might take advantage of aggregations of food items other than a nts. It is intriguing to speculate on the possibility that G. carolinensis might sequester natural products from ingested ants and mites, and use them to augment its integumentary defenses, much as some dendrobatid and ranid frogs do (Eisner et al. 1990, Saporito et al. 2012 and references therein). Developmental studies 2013 M. Deyrup, L. Deyrup, and J. Carrel 373 by Garton and Mushinsky (1979) show that the skin of G. carolinensis becomes intrinsically toxic and unpalatable to predaceous vertebrates as tadpoles metamorphose. However, to our knowledge, chemical analyses of organic products in skin secretions from this toad have not been performed. It is conceivable that as free-ranging toads feed they systemically retain substances of dietary origin and broaden their integumentary defenses, rendering them more effective against the diverse array of predators and parasites encountered in nature. If such chemical studies were to support the dietary-defense hypothesis, controlled feeding tests using taxonomically identified ants and mites could be warranted . Acknowledgments This research was supported by the Archbold Biological Station. We thank Nancy Deyrup for helping prepare Table 1. Literature Cited Abrahamson, W.G., A.F. Johnson, J.N. Layne, and P.A. Peroni. 1984. 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Ants Stomachs % total % non-ant Ants # % # % prey prey Ponerinae 17 0.35 Hypoponera 12 0.25 4 2.74 H. opaciceps (Mayr) 8 0.16 3 2.05 H. punctatissima (Roger)* 4 0.08 1 0.68 Odontomachus relictus Deyrup & Cover 5 0.10 5 3.42 Ecitoninae 197 4.05 Neivamyrmex 197 4.05 15 10.27 N. carolinensis (Emery) 84 1.73 3 2.05 N. opacithorax (Emery) 113 2.32 12 8.22 Myrmicinae 3410 70.03 Aphaenogaster 27 0.55 16 10.96 A. flemingi M.R. Smith 4 0.08 4 2.74 A. floridana M.R. Smith 1 0.02 1 0.68 A. miamiana Wheeler 2 0.04 2 1.37 A. treatae Forel 20 0.41 9 6.16 Cardiocondyla emeryi Forel* 1 0.02 1 0.68 Crematogaster ashmeadi Mayr 2 0.04 1 0.68 Cyphomyrmex 21 0.43 13 8.9 C. minutus Mayr 17 0.34 10 6.85 C. rimosus (Spinola)* 4 0.08 3 2.05 Monomorium viride Brown 39 0.80 3 2.05 Pheidole 2749 56.45 130 89.04 P. adrianoi Naves 10 0.21 4 2.74 P. dentata Mayr 12 0.25 3 2.05 P. floridana 522 10.72 56 38.36 P. metallescens Emery 399 8.19 39 26.71 P. moerens Wheeler* 371 7.62 14 9.59 P. morrisi Forel 1433 29.43 96 65.75 Pheidole, unidentified 2 0.04 2 1.37 Pyramica deyrupi Bolton 4 0.08 3 2.05 Solenopsis 475 9.76 75 51.37 S. abdita Thompson or carolinensis Forel 51 1.05 13 8.9 S. globularia littoralis Creighton 52 1.07 14 9.59 S. invicta Buren* 67 1.38 16 10.96 S. nickersoni Thompson 174 3.57 31 21.23 S. pergandei Forel 128 2.63 25 17.2 S. tennesseensis M.R. Smith 3 0.06 2 1.37 Strumigenys 21 0.43 4 2.75 S. emmae (Emery)* 3 0.06 2 1.37 S. louisianae Roger 1 0.02 1 0.68 S. rogeri Emery 17 0.35 1 0.68 Trachymyrmex septentrionalis (McCook) 71 1.46 27 18.49 Formicinae 1159 23.80 Brachymyrmex depilis Emery 149 3.06 21 14.38 Camponotus 2 0.04 2 1.37 C. floridanus (Buckley) 1 0.02 1 0.68 C. inaequalis Roger 1 0.02 1 0.68 2013 M. Deyrup, L. Deyrup, and J. Carrel 377 Ants Stomachs % total % non-ant Ants # % # % prey prey Nylanderia 1008 20.70 111 76.03 N. arenivaga Wheeler 433 8.89 81 55.48 N. concinna (Trager) 4 0.08 2 1.37 N. phantasma (Trager) 247 5.07 50 34.25 N. wojciki (Trager) 319 6.55 51 34.93 N. sp. 4 0.08 1 0.68 Dolichoderinae 76 1.56 Dorymyrmex 50 1.03 19 13.01 D. bossutus (Trager) 1 0.02 1 0.68 D. bureni (Trager) 30 0.62 7 4.79 D. elegans (Trager) 19 0.39 12 8.22 Forelius pruinosus (Roger) 26 0.53 1 0.68 Unidentified Formicidae 10 0.21 1 0.68 Formicidae, total 4869 100.00 146 100 95.35 Arthropods other than ants 266 101 69.18 5.18 Acarina 160 66 45.21 60.15 Arachnida 11 11 7.53 4.14 Coleoptera 32 23 15.7 12.03 Curculionidae Acalles minimus Blatchley 3 3 2.05 1.13 Scarabaeidae Aphodius campestris Blatchley 1 1 0.68 0.38 Ataenius sp. 1 1 0.68 0.38 Silvanidae Ahasverus rectus (LeConte) 5 3 2.05 1.88 Ciidae Cis sp. 5 5 3.42 1.88 Elateridae Conoderus bellus (Say) 3 3 2.05 1.13 Latridiidae Metophthalmus americanus Motschulsky 1 1 0.68 0.38 Tenebrionidae Poecilocrypticus formicophilus Gebien * 1 1 0.68 0.38 Staphylinidae Ischnosoma flavicolle (LeConte) 2 2 1.37 0.75 Proteininae sp. 1 1 0.68 0.38 Scydmaenidae sp. 4 4 2.74 1.5 Nitidulidae Stelidota geminata (Say) 5 1 0.68 1.88 Collembola 31 16 10.96 11.65 Entomobryidae sp. 3 3 2.05 1.13 Poduridae sp. 19 6 4.11 7.14 Sminthuridae sp. 9 8 5.48 3.38 Diptera 4 4 2.74 1.5 Ceratopogonidae sp. 4 4 2.74 1.5 378 Southeastern Naturalist Vol. 12, No. 2 Ants Stomachs % total % non-ant Ants # % # % prey prey Embioptera sp. 1 1 0.68 0.38 Heteroptera 3 3 2.05 1.13 Ceratocombidae Ceratocombus vagans McAtee & Malloch 1 1 0.68 0.38 Lygaeidae sp. 1 1 0.68 0.38 Cydnidae Melanaethus subpunctatus (Blatchley) 1 1 0.68 0.38 Hymenoptera (Non-ant) 5 5 3.42 1.88 Braconidae sp. 1 1 0.68 0.38 Encyrtidae sp. 1 1 0.68 0.38 Diapriidae sp. 1 1 0.68 0.38 Scelionidae sp. 1 1 0.68 0.38 Rhopalosomatidae Olixon banksii (Brues) 1 1 0.68 0.38 Orthoptera 2 2 1.37 0.75 Gryllidae Cycloptilum sp. 1 1 0.68 0.38 Tridactylidae Neotridactylus archboldi Deyrup & Eisner 1 1 0.68 0.38 Isoptera, Rhinotermitidae 16 5 3.42 6.02 Symphyla sp. 1 1 0.68 0.38