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Revista Chilena de Historia Natural 59: 201-208, 1986 Dietary specialization and variation in two mammalian myrmecophages (variation in mammalian myrmecophagy)

Especializaci6n dietaria y variaci6n en dos mamiferos mirmec6fagos (variaci6n en la mirmecofagia de mamiferos)

KENT H. REDFORD

Center for Latin American Studies, Grinter Hall, University of , Gainesville, Florida 32611, USA

ABSTRACT

This paper compares dietary variation in an opportunistic myrmecophage, novemcinctus, and an obligate myrmecophage, Myrmecophaga tridactyla. The diet of the common long-nosed , D. novemcintus, consists of a broad range of as well as and plant material. In the , and are less important as a food source than they are in . The diet of the giant . M. tridactyla, consists almost entirely of ants and termites. In some areas giant consume more ants whereas in others termites are a larger part of their diet. Much of the variation in the diet of these two myrmecophages can be explained by geographical and ecological variation in the abundance of prey. However, some variation may be due to individual differences as well. Key words: Dasypus novemcinctus, Myrmecophaga tridactyla, Tamandua, food habits. armadillo, , ants, termites.

RESUMEN

En este trabajo se compara la variacion dietaria entre un mirmecofago oportunista, Dasypus novemcinctus, y uno obligado, Myrmecophaga tridactyla. La dieta del armadillo comun, D. novemcinctus, incluye un amplio rango de in- vertebrados así como vertebrados y materia vegetal. En los Estados Unidos, hormigas y termites son menos importantes como recurso alimenticio de los , de lo que son en Sudamérica. La dieta del hormiguero gigante, M tridactyla, está compuesta casi enteramente por hormigas y termites. En algunas áreas el hormiguero gigante consume más hormigas que termites, en tanto que en otras las termites forman la parte más importante de su dieta. Mucho de la variaci6n dietaria en estos dos mirmecofagos puede explicarse por variaci6n geográfica y ecologica en la abundancia de presas. Sin embargo, alguna variaci6n puede deberse a diferencias individuales entre los mirmecofagos. Palabras claves: Dasypus novemcinctus, Myrmecophaga tridactyla, Tamandua, hábitos alirnenticios, armadillo, hormiguero gigante, hormigas, termites.

INTRODUCTION This comparison elucidates some of the factors governing dietary variation in inver- A great number and variety of ants and tebrate-eating . The dietary im­ termites provide food for a variety of portance of social as a group and of mammals. These and -eating ants versus termites varies for both of these mammals or myrmecophages are found on geographically, ecologically, and in- all tropical continents and include opportu- dividually. I will relate this variation to the nistic myrmecophages such as mongooses, differing availabilities of prey and the and obligate myrmecophages such as extent of myrmecophagy. (Redford in press a). In this paper I compare two South American mammals, the common long-nosed armadi- ANTS AND TERMITES AS FOOD llo, Dasypus novemcinctus, an opportu- nistic myrmecophage, and the giant an- In many parts of the world ants and teater, Myrmecophaga tridactyla, an obli- termites provide a large base of potential gate myrmecophage. prey for mammalian predators. Both ant

(Received 3 December 1985. Accepted 17 March 1986.) 202 REDFORD

and termite faunas are most diverse in contribution that ants make to invertebrate tropical latitudes with species diversity faunas increases with an increase in the dropping off sharply with increasing la- dryness of a habitat (Pisarski 1978). Ter- titude (Kusnezov 1957, Wilson 1971, Wood mites, on the other hand, have a primarily & Sands 1978, Mill 1982a). Single habitats tropical and sub-tropical distribution due in tropical have over 40 species of to their low mobility, their reliance on termites (Mill 1982b) whereas there are cellulose for food, and their susceptibility only 13 species in the entire state of to dessication. The termite species which Florida (Snyder 1954). The pattern is are found in temperate latitudes are fre- similar for ants with 455 species found in quently confined to the buffered environ- , north of Mexico (Wilson ment of dead wood (Wilson 1971, Wood & 1971 ), compared with 222 species known Sands 1978). from the Brazilian state of Sao Paulo alone When compared with other arthropods, (Kusnezov 1957). ants and termites are good food. Redford There are few data from the Neotropics and Dorea (1984) have shown that al- allowing quantification of the importance though lower in fat and higher in ash of ants and termites (Pisarski 1978). One content than some other , rough estimate provided by Fittkau & Klinge termites and ants have equivalent total ( 1973) based on their extensive work in the nitrogen contents. What makes these Amazon forest indicated that ants and insects particularly suitable as a source of termites composed over 30,. of the total food is their colonial habit and the con- biomass and about 75,. of the total sequent concentration of potential food. biomass. It is possible to compare this with the termite biomass calculated from three different studies. Lubin et al ( 1977) estimated that termites of the Nasutitermes had a biomass of 12.8 kg/ha MAMMALIAN MYRMECOPHAGES in a Panamanian forest. The biomass of all termites in the top five em of soil in an Amazonian forest averaged 4.3 kg/ha (Ban- Dasypus novemcinctus deira 1979) whereas in the Brazilian cerrado four species of termites averaged The long-nosed armadillo (Dasypus novem- 33.3 kg/ha (Redford 1984). cinctus), an opportunistic myrmecophage, For ants, the data are more scarce. is found not only in areas where ants and Levings & Franks (1982) and Levings & termites are common but also where they Windsor ( 1982), calculated an average of are scarce. The food habits of this species 663 ant colonies/ha in the Panamanian are well known in the United States be- rainforest and found that half of all of the cause armadillos were at one time sus- individual forest litter arthropods collected pected of eating the eggs of game birds. As over 30 months were ants. Based on many a result, the stomachs of hundreds of years of work in Peruvian forests, Erwin were painstakingly examined. Ta- (1983, p. 14) stated that: "It is also clear ble 1 summarizes the results of the three that ants constitute the greatest number of largest published studies of D. novem- individuals and biomass in the canopy as cinctus food habits in the United States. well as on the ground". This conclusion is In , where the largest study was supported by Adis et al. (1984) who found done (Kalmbach 1944) armadillos ate that ants accounted for between one- mostly and larvae. Other quarter and one-half of the total arthropod major foods included Orthoptera, centi- biomass obtained from the canopy of a pedes and millipedes, and Diptera. Hyme- Brazilian forest. noptera equalled 14.1.,. volumetrically, There are few data available to allow with the great majority being ants of 44 comparison of the relative abundances of different species. Termites of the genus ants and termites in single habitats. Ants Reticulitermes ocurred in 7 5,. of the are distributed over a much greater latitu- stomachs, although composing only 4.50fe dinal range than termites and unlike ter- by volume. Armadillos also consumed small mites, can be dominant elements in tem- amounts of earthworms, crayfish, carrion, perate arthropod faunas (Kusnezov 1957, , , small mammals, fruit Wilson 1971, Petal 1978) . The percent and mushrooms. DIETARY SPECIALIZATION AND VARIATION IN MYRMECOPHAGES 203

TABLE 1 South America where both ants and ter- mites are more common than in the United Relative composition of Dasypus novemcinctus States. The two available samples of D. diets, expressed as percent of total stomach volume novemcinctus stomachs from the southern Con;~ posicion relativa de la dieta de Dasypus novemcinctus, part of its range show a different picture expresada como porcentaje del volumen de la ingesta en el estomago from the results discussed before. Mathews ( 1977) analyzed two D. novemcinctus Florida stomachs from western Brazil. One of the Texas (n=104) (n =172) stomachs contained 90.,. termites of several (n=169) Fitch Nesbitt species with the remaining 10"fo containing Kalmbach et al. et al. Food Type 1944 1952 1977 ants, termites, beetles and scorpions. The second stomach contained a large quantity Hymenoptera 14.7 3.9 14.7 of what he guessed to be fruit with the rest Isoptera 4.7 1.2 0.1 consisting of mostly termites. Similar re- Coleoptera 43.5 44.6 29.0 Orthoptera 6.5 8.5 10.2 sults were encountered in a sample of five Lepidoptera 8.2 4.7 9 D. novemcinctus stomachs from central Other insects 2.6 5.8 13.5 Brazil (Redford unpubl. data). These Myriapoda 6.5 8.5 7.9 stomachs were found to contain an average Other invertebrates 8.3 8.5 11.0 Amphibia & Reptilia 1.3 4.5 1.8 of 69.,. ants and termites with the re- Other vertebrates 0.5 0 1.9 mainder beetles and millipedes. Another Plant material 2.2 9.8 0.8 five stomachs from the same locality ana- lyzed with a different methodology showed ants to comprise 17"fo by volume, termites 15"fo, and beetles and beetle larvae 41.,.. In Louisiana (Fitch et al. 1952), beetles One of the stomachs contained 65.,. fruit comprised an even larger proportion of the by volume, supporting Mathews ( 1977) diet with ants comprising only 3.9"fo, and observation. Several small vertebrates in- termites 1.2"fo. Plant material, including cluding a lizard and two skinks were seeds, berries and fungi comprised 9.8"fo. In also encountered. Florida (Nesbitt et al. 1977) approximately one third by volume of armadillos' diet was Other Dasypus beetles with ants comprising 15.1 "fo, and termites 0.1 "'"· Crayfish, small amphibians, The observation of increased consumption reptiles, and mammals such as nestling of ants and termites by D. novemcinctus in cotton rats (Sigmodon), were also con- South America is supported by examining sumed. the few data available for other species in In summary, in the United States D. the genus Dasypus. One D. kappleri novemcinctus is omnivorous, though prima- stomach from (Barreto et al. rily entomophagous, with beetles and their 1985) had 19.,. ants and 10.,. termites. One larvae comprising the largest single dietary D. sabanicola stomach from Colombia con- item. The percent of ants and termites in tained 10.,. ants and 88.,. termites (Barreto the diet varies from a low of 5.1.,. in et al 1985). Although a larger sample of Louisiana to a high of about 18.,. in Texas. the same species from the llanos of Vene- In all three samples ants were a much more zuela (Ferguson-Laguna 1984) contained important food than termites. Despite the mostly beetles and only 7.3.,. ants and fairly low representation of ants and ter- termites, other D. sabanicola stomachs mites, all authors have commented that, from (Pacheco & Naranjo 1978) when available, ants and or termites com- had 45% termites, 22"fo ants and only 18"fo prise one of the favored foods of D. beetles. Finally, one D. hybridus stomach novemcinctus and many of the stomachs from (Barlow 1965) contained examined contained thousands of ants and 49.,. ants and 9"fo termites, other assorted their coccons or termites and their nymphs. invertebrates and the remains of several In fact in some smaller samples (Newmann nestling mice. 1913) ants were said to be the most In summary, it is clear that despite small common prey encountered in the sample sizes there is a strong suggestion of stomachs. geographical variation in the consumption Dasypus novemcinctus unfortunately, of ants and termites both within D. novem- has been much less studied in Central and cinctus, and within the genus Dasypus as a 204 REDFORD whole (Fig. I). The general pattern appears young mice offered to captive giant an- to be a greater consumption of both ants teaters were consumed with great relish and termites in tropical areas. More ants (Redford 1983). than termites are consumed in temperate areas such as the United States and Uru- Tamandua guay, and in areas which are otherwise unsuitable for terrestrial termites such as The genus Myrmecophaga is monotypic so the seasonally flooded llanos of Venezuela. congeneric comparisons are impossible, but food habits of two species in a closely Myrmecophaga tridactyla related genus, Tamandua, may be examined. As with the giant anteater, tamanduas show The second edentate is the giant anteater, geographical variation in diet with some Myrmecophaga tridactyla, an obligate stomachs reported to contain only ants and myrmecophage. Although the data consist others only termites (Redford 1983). Indi- of few samples, this species appears to vidual differences in feeding by Tamandua consume virtually nothing but ants and contribute to variation in patterns of termites. However, as in the long-nosed myrmecophagy (Montgomery & Lubin armadillo, there is geographical variation in 1977). For example, two T. tetradactyla the diet (Table 2, Redford 1985). In the from Venezuela consistently spent very Venezuelan llanos ants comprise virtually different amounts of time foraging for the entire diet whereas in northern Brazil termites at nests versus in wood (Lubin & the entire diet seems to consist of termites. Montgomery 1981 ). They also demons- Within a much smaller spectrum of prey trated differences in feeding bouts with one types it appears that the giant anteater, like animal spending 901ft on termites and the the long-nosed armadillo, is also an oppor- other only 241ft (Lubin & Montgomery tunist. unpubl.). Similarly, two T. mexicana indi- viduals fed for different amounts of time on TABLE 2 the same prey types (Montgomery & Lubin 1977). Food habits of Myrmecophaga tridactyla based As in the giant anteater, anteaters of the on stomach contents, feces and observations (from genus Tamandua are capable of eating prey Redford 1985) other than ants and termites. One zoo-re- Habitos alimenticios de Myrmecophaga tridactyla, basado searcher (Parker pers. comm.) reported en contenidos estomacales, fecas y observaciones directas tamanduas killing and consuming iguanine (tornado de Redford 1985) lizards housed in the same cage.

Location 1ft Ants 1ft Termites DISCUSSION 100 0 Ceara, Brazil 100 0 Both Dasypus novemcinctus and Myrmeco- N.E. Brazil 100 0 phaga tridactyla have large geographical Llanos of Venezuela 99 1 ranges within which they are found in Minas Gerais, Brazil 88 12 Brasilia, Brazil 15 85 extremely diverse habitats ranging from the Goias, Brazil 11 89 dry Paraguayan chaco to the wet forests of Amapa, Brazil 0 100 Amazonia (Fig. 2). Dasypus novemcinctus is found both farther north and farther south than the giant anteater; the latter's more limited range may reflect the prima- There have not been any detailed studies rily tropical distribution of extensive popu- on variation in the diet of Myrmecophaga, lations of ants and termites which form its but this species will take prey other than major diet. The long-nosed armadillo's ants and termites. One stomach from Boli- range is apparently limited by cold tempe- via contained only beetle larvae (Krieg ratures which affect the availability of soil 1939), whereas examination of feces from and litter invertebrates (Humphrey 1974, southern Brazil showed consumption of McNab 1980). millipedes (Shaw et al. In press) Anteaters These two edentates have diets with very in a U.S. zoo consumed nestling cottontail different size distributions. The armadillo rabbits (MacNamara pers. comm.), and does not specialize on particular classes, DIETARY SPECIALIZATION AND VARIATION IN MYRMECOPHAGES 205

1 2

Q.!!:_ Florida% volume

D .n. Mato Grosso, Brazil (n= 1) D.n. Goias, Brazil (n :5:s.s.)

!2_, hybridus (n = 1)

Fig. 1. Summary of the relative consumption of ants and termites by four species of Dasypus in various parts of North and South America. (D.n. = Dasypus novemcinctus). The circles represent 100"1o of the diet, the wedge numbered "1" equals the relative percent of the diet composed of ants, the wedge numbered "2" equals the relative percent composed of termites. The sources of the data are listed in the text. Resumen del consumo relativo de hormigas y termites por parte de cuatro especies de Dasypus en varias partes del Norte y Sudamerica. (D.n. =Dasypus novemcictus). Los cfrculos representan lOO"'o de Ia dieta; Ia cuiia numerada "1" representa el porcentaje de Ia dieta que esta compuesto por hormigas; Ia cuiia numerada "2" representa el porcentaje de Ia dieta que esta compuesto por termites. Las fuentes bibliograficas de los datos estan listadas en el texto. 206 REDFORD

taking very small prey as well as compara- prey that can be ingested. The proportion tively large prey. A long-nosed armadillo of ants versus termites in the diet probably stomach may be thought of as a Berlese reflects the relative availability of each to funnel, reflecting, with some biases, the the anteater and is not based on factors composition of the leaf litter fauna. In intrinsic to particular prey species. This South America, where ants and termites are would allow the giant anteater to range more important components of the ground throughout many different habitats with dwelling invertebrate fauna, they make up their different tropical social insect com- a greater percentage of its diet. However, munities (Redford 1985). throughout its range this species will eat Both Dasypus novemcinctus and Myrme- small vertebrates and fruit when available. cophaga tridactyla respond opportu- Dasypus novemcinctus' extensive range re- nistically to local availabilities of prey. This flects a virtually unprecedented ability to opportunism by long-nosed armadillos is utilize whatever foods are locally available illustrated by seasonal trends in prey con- (Redford in press b). sumption. In the U.S. some prey such as Orthoptera, reptiles and amphibians are more commonly taken in the colder months when low temperatures slow them down (Kalmbach 1944, Fitch et a/. 1952, Talmage & Buchanan 1954). Diptera and Lepidoptera larvae are taken in the spring and summer when plentiful (Kalmbach 1944, Fitch et a/. 1952) whereas earth- are most common in the diet during the wet season when a high water table forces them up (Kalmbach 1944, Nesbitt et a/. 1977). The consumption of ants and termites by Dasypus is probably also affected by seasonal factors. In the Florida data (Nes- bitt eta/. unpubl.), the trend is for ants to be taken more frequently in the wet season. This also occurs in the Venezuelan llanos (Ferguson-Laguna 1984). Termites are very responsive to humidity levels and are much more common above ground during the wet season. Therefore, in areas where termites are common, one would expect them to be a more common prey item for surface-foraging armadillos during the rainy season. The giant anteater also responds to local variation in prey availability. As is the case Fig. 2. The geographical ranges of the common with D. novemcinctus some of this res- long:nosed armadillo. Dasypus novemcinctus and ponse can be explained by correlations of the giant anteater, Myrmecophaga tridactyla. prey abundance with humidity and rainfall. Los rangos geogrwcos del armadillo comun, Dasypus Ants were reported to compose I 00% of novemcinctus, y del hormiguero gigante, Myrmecophaga the diet in very dry areas where termites tridactylll. would be expected to be rare. The other area where ants comprise a high percentage of the diet is the seasonally flooded V ene- The giant anteater on the other hand, zuelan llanos which is unsuitable for terres- consumes mostly small invertebrates, trial termites. specializing on small arthropods found in In conclusion, both Dasypus novem- dense aggregations (i.e., ants and termites). cinctus and Myrmecophaga tridactyla are This strict myrmecophagy is reflected by opportunistic predators, responding to lo- the highly specialized of the giant cal availabilities of prey. Their responses anteater which severely limits the type of are in large part shaped by the degree to DIETARY SPECIALIZATION AND VARIATION IN MYRMECOPHAGES 207 which they have specialized on ants and KUSNEZOV N (1957) Numbers of species of ants in termites. The available data, summarized in faunae of different latitudes. Evolution 11: 298-299. this paper, allow formulation of four hypo- LEVINGS SC &. DM WINDSOR (1982) Seasonal and theses. First, Dasypus armadillos consume a annual variation in litter arthropod populations. particular prey type in proportion to its In: Leigh EG Jr, AS Rand&. DM Windsor (eds) availability. Second, over much of the The ecology of a tropical forest 355-387, Smithsonian Press, Washington DC. tropical portion of their range the diets of LEVINGS SC &. NR FRANKS (1982) Patterns of nest Dasypus armadillos will consist largely of dispersion in a tropical ground ant community. ants and termites. Third, in any given Ecology 63: 338-344. habitat Myrmecophaga will prefer the most LUBIN YD &. GG MONTGOMERY (1981) Defenses of available species of ant or termite. Fourth, Nasutitermes termites (lsoptera, Termitidae) against Tamandua anteaters (Edentata, Myrme- individual variation accounts for a signi- cophagidae). Biotropica 13: 66-76. ficant proportion of the within-population LUBIN YD, GG MONTGOMERY &. OP YOUNG (1977) variation in feeding by anteaters. Food resources of anteaters (Edentata: Myrme- cophagidae) I. A year's census of arboreal nests of ants and termites on Barro Colorado Island, Panama Canal Zone. Biotropica 9: 26-34. ACKNOWLEDGMENTS MATHEWS AGA (1977) Studies on termites from the I would like to thank J.F. Eisenberg, P.L. Meserve, J.G. Mato Grosso State, Brazil. Academia Brasileira Robinson, P. Shaw and two reviewers for help with this de Ciencias, Rio de Janeiro. paper. The Florida Game and Fresh Water Fish Commis- MCNAB BK (1980) Energetics and the limits to a sion provided valuable access to unpublished data; Mrs temperate distribution in armadillos. Journal of WM Hetrick helped in the analysis of the armadillo Mammalogy 61: 606-627. stomachs; and P. Parker, M. MacNamara, and Y.D. Lubin MILL AE (1982a) Foraging and defensive behaviour in provided unpublished information. neotropical termites. PhD dissertation, Univer- sity of Southampton. 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