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Research Question: How does species size affect the eating habits of anurans?

Predictions: We hypothesize that size does matter. The diets of larger species of frogs will be more diverse than those of smaller species.

Purpose: We hope to learn more about the ecological niches of frogs, and the environmental implications therein. Obviously, this project cannot provide a complete explanation of frogsŐ niches. We will not examine what predators eat frogs, or what exact habitats different frog species live in. We cannot even offer the frogs every single food option they might have in the wild; a few very different types of food will have to suffice. However, this project can potentially give us some general insights into how much variety the species we will study are adapted to handle. This, in turn, may be relevant to some important environmental issues regarding the habitats of these frogs and their food (see "Relevance of Research").

General Information: Anurans (frogs and toads) are amphibians, or small vertebrates that undergo partial metamorphosis, and spend part of their lives in water, and part on land. Amphibian eggs are gelatinous, and are laid in water. Young anurans hatch from the eggs as tadpoles, purely aquatic, and then mature into air-breathing adults with legs. Frogs will remain around bodies of water even as adults. Toads will leave the water as soon as they reach adulthood, and not come back except to mate and lay eggs.

Also, amphibians are ectothermic (cold-blooded), which means that their bodies are the same temperature as the environment immediately around them. In the winter, amphibians in temperate climates will hibernate, by burying themselves as deeply as possible, and becoming comatose for the duration of the season. Some frogs do not need to insulate themselves by burying in the ground. Instead, they simply freeze solid, and revive when they are thawed by warmer weather.

Anurans typically eat small invertebrates (insects, small spiders, worms, etc.), although some big frogs are also knownto eat small vertebrates, such as fish or smaller frogs. By doing this, anurans fill a role in the ecosystem, called a niche. If different species eat different types or varieties of invertebrates and small vertebrates, then they fill slightly different specific niches

Relevance of Research: Do species of different sizes compete for the same food, or are they filling completely separate niches? Is size an adaptation to allow frog species to eat certain selections of foods? One implication of this could be that frog species with very limited diets would be more susceptible to loss of prey species. If small frog species do have diets of limited variety, as predicted, then their populations might decline quickly when prey species start to disappear. Conversely, species capable of eating a variety of foods (the larger species, if our hypothesis is correct) would not suffer very much from the loss of one or two prey species. If a large frog cannot find insects, it might survive easily on earthworms, fish, and even smaller frogs. A smaller frog would not have that luxury. In this scenario, the small frogs would be very sensitive environmental indicator species. So, if our hypothesis is correct, we could proceed in future studies to look at where small frog species are disappearing. If Blanchard's Cricket Frogs (Acris crepitans blanchardi), which are very small, are not equipped to eat fish or other aquatic foods, would they disappear from wetlands with few land insects? According to some literature, "large areas of terrestrial habitat surrounding wetlands are critical for maintaining biodiversity" (Semlitsch, p. 1219). Could cricket frogs show the symptoms of inadequate terrestrial buffers (such as a riparian corridor) between wetlands and human development?

When frogs do compete for the same food, does size give some species advantages over others? This may be the case with some invasive species. American Bullfrogs (Rana catesbeiana), which have been introduced to the western United States from other portions of the country, are reportedly out-competing the local, smaller frogs, and other small predators. "On Buenos Aires and nearby San Bernadino refuge, bullfrogs nearly eliminated the Mexican garter snake and completely wiped out the threatened Chiricahua leopard frog, species that were common on both refuges only decades ago." Meanwhile, the invading bullfrogs are becoming overpopulated, with population densities "20 to 30 times higher than they are back East" (Tangley, p. 42). Understanding the affect of size on ability to catch various foods may reveal possible reasons for the bullfrog's success. Does the bullfrog thrive because it eats more variety of food than the leopard frog, or does it eat the same food in more quantity? Or does the bullfrog skip competition altogether and simply eat the smaller leopard frogs?

Literature Review: The following references are examples of current literature that is related to this project and its various implications. It must be noted that none of this literature directly addresses this exact experiment, because (to the best of our knowledge and research), there are no available records of studies on the correlation between size and diet in anurans.

Anonymous. "Care Sheets." Available: http://www.amphibiancare.com/frogs/caresheets.html

This webpage provides identification information on species of frogs and toads, along with instructions on how to care for each species in captivity. This might help us to properly keep our specimens, without which the project would not rock. The site shows specifics, such as aquarium tank sizes and water temperature, that are optimum for each of the frog species.

Alvarez, D.; Nicieza A. G. "The Effects of Temperature and Food Quality on Anuran Larval Growth and Metamorphosis". Functional Ecology, Vol. 16 (5): pp. 640-648.

This article's focus on food quality gives an idea of why anurans eat what they eat.

Arendt, J. D. "Reduced burst speed is a cost of rapid growth in anuran tadpoles: problems of autocorrelation and inferences about growth rates." Functional Ecology. Vol. 17 (3), pp. 328-334.

This article shows how size and growth affect swimming speed in tadpoles. Rapid growth and large size result in slower speed. This may also be true in adult frogs. If so, it would work against the ability of large frogs to catch moving prey, such as crickets. However, our specimens will be kept in relatively small areas, with plenty of food, so catching food will not be a big problem. If the larger frogs still seem to prefer slow-moving food, such as worms, that might imply an biological adaptation to compensate for sluggishness. If there is no such adaptation, this phenomenon might not show up at all, unless the frogs become extremely fat.

Brodman, Robert; Cortwright, Spencer; Resetar, Alan. "Historical Changes of Reptiles and Amphibians of Northwest Indiana Fish and Wildlife Properties." The American Midland Naturalist. Vol. 147 (1), pp. 135-144.

This article explains that large scale habitat loss and change has affected the numbers and distribution of anurans in some areas. It provides an insight into the affect of alteration of land use on food availability and frog diversity.

Cogalniceanu, Dan; Palmer, Michael; Cibuc, Constantin. "Feeding in Anuran Communities on Islands in the Danube Floodplain". Amphibia-Reptilia, Vol. 22 (1): pp. 1-19.

This article states that, in their experiment, "Overall species with a larger size range consumed a higher prey diversity," which very directly supports our hypothesis. However, this article studies species other than those in our experiment, which leaves us with a relatively unanswered question: Does this apply to Ohio anurans as well?

Davis, Jeffrey G.; Menze, Scott A. In Ohio's Backyard: Frogs and Toads. Columbus: The Ohio Biological Survey, 2002.

This book provides a wealth of fantastic information about all of the fourteen species of frogs and toads in Ohio.

"Feeding of Casualty Amphibians" UK Wildlife Casualty Management. Available: http://212.187.155.84/wnv/Subdirectories_for_Search/Health&Management_Contents/UKFirstAidandCare/IndividualFeeding/Cas_Feeding_Amphibians.htm

This is an important article to us as it explains how we can go about feeding the frogs, what precautions to take, what food they will readily eat, and how to present it. It also points out that frogs often may refuse to eat in captivity.

"Frogs: An Early Indicator of Global Disaster." Solcomhouse. Available: http://www.solcomhouse.com/frogs.htm

This website, similar to Frog Web, provides information on environmental problem affecting anurans, and on the various specific effects of those problems.

"FrogWeb: Amphibian Declines & Deformities." National Biological Information Infrastructure. Available: http://www.frogweb.gov/

This website shows the frog decline "hotspots" in the US, such as California, and worldwide as well. It offers a wealth of information on the plights of threatened anuran species across the globe. If we end up wanting to use the results of this project to make predictions of what diet-related problems might affect frog populations, this website could likely help us find out where such problems may be occurring.

Hirai, Toshiaki. "Ontogenetic Change in the Diet of the Pond Frog, Rana nigromaculata". Ecological Research, Vol. 17 (6): pp. 639-644.

This article states that "as frogs increased in size, they increased total diet volume, but decreased the number of prey items." This seems to contradict our hypothesis, but once again, the study uses a different species than we will. Also, this study tracks the diet of one species at different ages and sizes of the individual specimen; our project will compare the diets of differently-sized species, dealing primarily with average species size instead of specimen size.

Hirai, Toshiaki; Matsui, Masafumi. "Feeding Habits of the Japanese Tree frog, Hyla japonica, in the Reproductive Season." Zoological Science, Vol. 17 (7), pp. 977-982.

This article shows that some frog species are opportunistic feeders, hence a varied diet. It raises questions about what impact this has on their distribution: Are the opportunistic frogs more widely distributed and are they more successful?

Marsh, David M.; Goicochea, Marco A. "Monitoring Terrestrial Salamanders: Biases Caused by Intense Sampling and Choice of Cover Objects." Journal of Herpetology, Vol. 37 (3), pp. 460.

This article examines the problems with some types of testing equipment, such as wooden artificial habitats for salamanders, and how they can skew experiment results. This raises the question of how the habitat we give our frogs may affect their behavior, and hence affect our desired results.

Matsui, Masafumi; Hirai, Toshiaki. "Food Habits of an Endangered Japanese Frog, Rana porosa brevipoda". Ecological Research, Vol. 16 (4): pp. 737-743.

This article examines a frog with a very diverse diet, which could be compared to the diets of our specimens.

"Missouri's Toads and Frogs." Missouri Conservation Commission (Reprinted from the Missouri Conservationist, by Tom R. Johnson). Available: http://www.conservation.state.mo.us/nathis/herpetol/frog/frog2.htm

This is basically a guide book that will help us with identification of the different species we are likely to come across here. It details species information, such as size range, diet, coloration, habitat, geographic range, some general behavior, and more.

Semlitsch, Raymond D.; Bodie, J. Russell. "Biological Criteria for Buffer Zones around Wetlands and Riparian Habitats for Amphibians and Reptiles." Conservation Biology, Vol. 17 (5), pp. 1219.

This article gives as an insight to the importance of terrestrial buffer zones and their impact on semi-aquatic organisms, such as amphibians. Frogs that require primarily terrestrial invertebrates as food also require undisturbed terrestrial habitat around their wetland habitat, in which to find that food. This would be particularly true for frogs that eat a limited variety of foods, since having the habitat for their primary foods available would be more critical.

Somsueb, P; Boonyaratpalin, M. "Optimum protein and energy levels for the Thai native frog, Rana rugulosa weigmann." Aquaculture Research, Vol. 32 (12), pp. 33-38.

This article shows that frogs have particular nutritional requirements, which are met by the types of food in their diet. Frogs that can get the nutrition that they need from particular foods are prey-specific in their natural habitats.

Stocum, David L. "Frog limb deformities: An 'eco-devo' riddle wrapped in multiple hypotheses surrounded by insufficient data." Teratology, Vol. 62 (3), pp. 147-150.

This article serves as a warning to keep the experiment simple and straightforward. It is an example of what not to do: if the project gets too complex and confused, trying to make sense of the results can get ugly.

Tangley, Laura. "A Plague of Aliens." National Wildlife, Vol. 41 (2), pp. 42-45.

This article discusses the environmental problems caused by several invasive species in the United States, focusing greatly on the bullfrog, which "consumes virtually any living thing it can fit into its enormous mouth." Since the bullfrog is very large, this supports our hypothesis that larger frogs eat a greater variety. More importantly, it addresses one major possible implication of this project: frogs that can have more varied diets are more able to adapt to changes in habitat. This is not always good, because that ability allows such species to wreak havoc if they are introduced to habitats in which they do not belong.

Overview: We will offer a variety of food options to individual specimens and observe which they choose to eat. Data will be measured in two ways: first, in numbers of each food item eaten by each specimen; and second, in biomass gained by each specimen over time. We will keep all of the frogs in separate areas, so as to be sure which specimen eats which food item. We will offer the same food types to all specimens. However, the size of the food may have to vary in accordance with the size of the specimen, as small frogs simply cannot swallow very large food, while large frogs my ignore very small food.

The Specimens: We will experiment with three species of anurans: Northern Green Frogs (Rana clamitans melanota), Blanchard's Cricket Frogs (Acris crepitans blanchardi), and Eastern American Toads (Bufo americanus americanus). We will use five specimens from each species, in order to help provide average data for each species, and avoid irregularities that might come from an eccentric individual specimen. More than five specimens per species would provide more significant averages, but larger numbers of specimens are beyond the capabilities of our resources for this project.

The Food: We will offer crickets, red wiggler worms, meal worm larvae, and guppies (small fish) as food options for the anurans. Every specimen will get all four options at every feeding. We will attempt to make the size of the food organisms appropriate for the size of the specimen. (No giving full-grown crickets to the cricket frogs; the specimen must be somewhat bigger than the food.) We will also attempt to make quantities of food approximately appropriate, yet abundant, for the specimens. There should not be massive amounts of food left over, but no specimen's feeding should be limited by the amount it is given.

Materials: Specimens will be kept in five ten-gallon glass aquariums. Each tank (aquarium) will be divided into three sections by plastic dividers, in order to keep the individual specimens and their food separated. Each section will be outfitted with an area of "land," in the form of dry rocks (no soil, so that uneaten food will always be visible to the observer; we do not want worms to bury themselves), and an area of clear water.

Crickets will be kept in a large plastic trash can, with a lid (kept on only loosely, so that the crickets have air). We will attempt to keep smaller crickets in a second trash can. We cannot breed crickets, so we will obtain more when necessary. Red wiggler worms will be kept and bred in a vermicompost worm bin, and fed organic compost. Meal worms will be bred in another bin, in a bed of oats and wheat meal, and fed raw potato or apple slices. Only the larval form will be fed to the specimens (the adult is a beetle). Guppies will be bred in a fish tank, and fed fish food. We have been advised to place small decorative objects in this tank to keep the fish cheerful.

A plastic cup will be used to carry crickets, worms, and meal worms to the anurans. A fish net will be used for the fish. When a specimen is weighed, it will be caught with a net and placed in an empty plastic container, which will be placed on a scale.

Methods of Feeding: Group members will take turns tending to the experiment. This includes feeding food populations (to be done as needed), recording data, and feeding specimens. Specimens will be fed every other day. When specimens are fed, live leftovers from the previous feeding will remain in the tanks. Dead leftovers will be removed. If a specimen has eaten all of the food from one type that it has been given, it should receive a larger quantity of food of that type. (If a toad wants to eat twenty meal worms, it should be able to do so.) If a specimen shuns a food option completely, however, the option must still be offered, just in case the specimen changes its mind.

Methods of Measuring: Each time the specimens are fed, the cages must first be checked for uneaten food from the previous feeding. Numbers and types of leftovers must be recorded for each specimen at each feeding. Whether the leftovers are dead or alive does not matter for this. Also, the amount of each type of food given to each specimen at the feeding time must be recorded. Numbers of leftovers can be subtracted from numbers of food organisms given at the previous feeding to calculate the number of each food that was eaten between feedings.

Once per week, all specimens must be weighed and the weights recorded. Specimens will be weighed one at a time. They can be placed in a container on the scale, so that they do not jump away. The weight of the container when empty can be subtracted from the weight of the specimen-and-container combination to get the weight of the specimen.

Data Tables: Below are blank data sheets of the formats that will be used for numerical data and weight data, respectively:

Timeline for Research: The specimens will be weighed and fed for the first time as soon as all specimens have been obtained, and all tanks set up. This will likely be on or before October 16, 2003. After that point the routine described above (see "Methods of Feeding" and "Methods of Measuring") will be observed until the completion of the project at the end of the semester.

Observations: Anecdotal observations will be noted in writing, and through pictures and video clips. These will all show up in progress reports, and in the final report. Possible explanations for phenomena in the results will also be included (for example, maybe toads do not like fish because adult toads are not adapted for water).

Results Tables and Graphs: Below is a blank table for numerical data averages for each specimen and each species. We will also create a line chart based upon the weight of the specimens over time, with a separate line for each specimen, and a second line chart for the average weight of the species over time. We do not have blank line charts created yet. In addition to these displays, we will use T-Tests to determine if the differences between species in average food type numbers are significant, and also to see if the differences between species in average weight gain are significant. We may also research other statistical methods to test for a correlation between average species diet and average species weight.

In the final paper, the results, implications thereof, and applications to bigger issues will be thoroughly discussed. Will the hypothesis be supported? Do bigger anurans eat a larger variety of foods? We will find out. Stay tuned. . . .

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• Frog Pictures Test (10/24/03) [ Contributor: Leo Sack ]
• From Hays: Frog Cricketville (10/16/03) [ Contributor: Hays Cummins ]
• A message from your bro' (10/15/03) [ Contributor: Carl M. Sack, Northland College ]

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