Variations in total brain mass and in the mass of three brain regions (main olfactory bulb, hippocampus, auditory nuclei) were examined using a data set for 63 species of bats (Chiroptera). Using both conventional and phylogenetically based analysis of covariance (log body mass as covariate), we tested several hypotheses that relate total brain mass or the size of the components to variation in foraging ecology, categorized as phytophagous, gleaner, and aerial insectivore. In some analyses, the category phytophagous was split into phytophagous pteropodid and phytophagous phyllostomid to examine differences between two distinct clades of bats. Because the Megachiroptera orient primarily by vision and olfaction, whereas all other bats rely on laryngeal echolocation to locate their prey, we hypothesized that the former would differ in size of the main olfactory bulb, as compared with all other bats. This hypothesis was supported by our analyses. Our more general prediction was that insectivorous bats, which rely heavily on echolocation for the pursuit and capture of their prey, would have larger auditory nuclei than do phytophagous species. This, too, was supported. We also compared phytophagous (fruit or nectar consuming) bats in two families, the Pteropodidae and the Phyllostomidae. We hypothesized that the phyllostomids, which use echolocation while foraging, would have larger auditory nuclei. Although statistical power is low in phylogenetically informed comparisons of the two clades, we did find weak evidence in support of this hypothesis. We conclude that bat brains show evidence of adaptation to foraging ecology.

1.
Altringham, J.D. (1996) Bats: Biology and Behavior. Oxford University Press, Oxford, UK.
2.
Arita, H.T., and M.B. Fenton (1997) Flight and echolocation in the ecology and evolution of bats. Trends Ecol. Evol., 12: 53–58.
3.
Barclay, M.R., and R. Brigham (1991) Prey detection, dietary niche breadth, and body size in bats: why are aerial insectivorous bats so small? Am. Nat., 37: 693–703.
4.
Baron, G., and P. Jolicoeur (1980) Brain structure in Chiroptera: Some multivariate trends. Evolution, 34: 386–393.
5.
Baron, G., H. Stephan, and H.D. Frahm (1996) Comparative neurobiology. In Chiroptera. 3 Volumes. Birkhäuser, Basel.
6.
Barton, R. (1999) The evolutionary ecology of the primate brain. In Comparative Primate Socioecology (ed. by P.C. Lee), Cambridge University Press: Cambridge.
7.
Barton, R.A., and P.H. Harvey (2000) Mosaic evolution of brain structure in mammals. Nature, 405: 1055–1058.
8.
Barton, R.A., A. Purvis, and P.H. Harvey (1995) Evolutionary radiation of visual and olfactory brain systems in primates, bats, and insectivores. Phil. Trans. R. Soc. Lond. B, 348: 381–392.
9.
Brashares, J., T. Garland, Jr., and P. Arcese (2000) The ecology, behavior, and phylogeny of the African antelope. Behav. Ecol., 11: 452–463.
10.
Clancy, B., R.B. Darlington, and B.L. Finlay (1999) Translating developmental time across mammalian species. Neuroscience, 105: 7–17.
11.
Cox, P. A., T. Elmqvist, E. D.Pierson, and W.E. Rainey (1991) Flying foxes as strong interactors in South Pacific island ecosystems: A conservation hypothesis. Conserv. Biol. 5: 448–454.
12.
Cruz-Neto, A.P., T. Garland, Jr., and A.S. Abe (2001) Diet, phylogeny, and basal metabolic rate in phyllostomid bats. Zoology, 104: 49–58.
13.
de Queiroz, K. (1996) Including the characters of interest during tree reconstruction and the problems of circularity and bias in studies of character evolution. Am. Nat., 148: 700–708.
14.
de Winter, W., and C.E. Oxnard (2001) Evolutionary radiations and convergences in the structural organization of mammalian brains. Nature, 409: 710–714.
15.
Deaner, R.O., C.L. Nunn, and C. P. van Schaik (2000) Comparative tests of primate cognition: Different scaling methods produce different results. Brain Behav. Evol., 55: 44–52.
16.
Devoogd, T.J., J.R. Krebs, S.D.Healy, and A. Purvis (1993) Relations between song repertoire size and the volume of brain nuclei related to song: comparative analyses amongst some oscine birds. Proc. R. Soc. Lond. B, 254: 75–82.
17.
Diaz-Uriarte, R. and T. Garland, Jr. (1998) Effects of branch length errors on the performance of phylogenetically independent contrasts. Syst. Biol., 47: 654–672.
18.
Felsenstein, J. (1985) Phylogenies and the comparative method. Am. Nat., 125: 1–15.
19.
Fenton, M.B. (1984) Echolocation: implications for ecology and evolution of bats. Q. Rev. Biol., 59:33–53.
20.
Fenton, M.B., A. Audet, M.K. Obrist, and J. Rydell (1995) Signal strength, timing, and self-deafening: The evolution of echolocation in bats. Paleobiology, 21: 229–242.
21.
Finlay, B.L., and R.B. Darlington (1995) Linked regularities in the development and evolution of mammalian brains. Science, 268: 1578–1584.
22.
Fleming, T.H., E.R. Heithaus, and W.B. Sawyer (1977) An experimental analysis of the food location behavior of frugivorous bats. Ecology, 58: 619–627.
23.
Freeman, P.W. (1981) A multivariate study of the family Molossidae (Mammalia: Chiroptera): morphology, ecology, and evolution. Fieldiana Zool., n.s., no. 7.
24.
Garland, T., Jr., A.W. Dickerman, C.M. Janis, and J.A. Jones (1993) Phylogenetic analysis of covariance by computer simulation. Syst. Biol., 42:265–292.
25.
Garland, T., Jr., P.H. Harvey, and A.R. Ives (1992) Procedures for the analysis of comparative data using phylogenetically independent contrasts. Syst. Biol., 41: 18–32.
26.
Garland, T., Jr., P.E. Midford, and A.R. Ives (1999) An introduction to phylogenetically based statistical methods, with a new method confidence intervals on ancestral values. Am. Zool., 39: 374–388.
27.
Griffiths, T.A. (1994) Phylogenetic systematics of slit-faced bats (Chiroptera, Nycteridae) based on hyoid and other morphology. Am. Mus. Nov., 3090: 1–17.
28.
Griffiths, T.A., A. Truckenbrod, and P.J. Sponholtz (1992) Systematics of Megadermatic bats (Chiroptera, Megadermatidae), based on hyoid morphology. Am. Mus. Nov., 3041: 1–21.
29.
Hand, S.J., and J.A.W. Kirsch (1998) A southern origin for the Hipposideridae (Microchiroptera)? Evidence from the Australian fossil record. In: Bat Phylogeny, Morphology, Echolocation and Conservation Biology (ed. by T.H. Kunz and P.A. Racey), Smithsonian Institution Press, Washington, D.C.
30.
Healy, S., and T. Guilford (1990) Olfactory bulb size and nocturnality in birds. Evol. 44: 339–346.
31.
Hutcheon, J.M. (2001) Molecular Systematics of the Chiroptera: A Phylogenetic Analysis Using DNA-DNA Hybridization and the Comparative Method. PhD dissertation, University of Wisconsin, Madison, WI.
32.
Hutcheon, J.M., J.A.W. Kirsch, and J.D. Pettigrew (1998) Base compositional biases and the bat problem. III. The question of microchiropteran monophyly. Phil. Trans. R. Soc. Lond. B, 353:607–617.
33.
Iwaniuk, A.N., and J. E. Nelson (2001) A comparative analysis of relative brain size in waterfowl (Anseriformes). Brain Behav. Evol., 57: 87–97.
34.
Jarrard, L.E. (1995) What does the hippocampus really do? Behav. Brain Res., 71: 1–10.
35.
Jolicoeur, P., and G. Baron (1980) Brain center correlations among Chiroptera. Brain Behav. Evol., 17: 419–431.
36.
Kirsch, J.A.W., T.F. Flannery, M.S. Springer, and F.-J. Lapointe (1995) Phylogeny of the Pteropodidae (Mammalia: Chiroptera) based on DNA hybridization, with evidence for bat monophyly. Aust. J. Zool., 43: 395–428.
37.
Kirsch, J. A. W., J.M. Hutcheon, D.G.P. Byrnes, and B.D. Lloyd (1998) Affinities and historical zoogeography of the New Zealand short-tailed bat, Mystacina tuberculata Gray 1843, inferred from DNA-hybridization comparisons. J. Mamm. Evol., 5: 33–64.
38.
Knudsen, J.T. and L. Tollsten (1995) Floral scent in bat-pollinated plants: a case of convergent evolution. Bot. J. Linn. Soc., 119: 45–57.
39.
Krebs, J.R., N.S. Clayton, S.D. Healy, D.A. Cristol, S.J. Patel, and A.R. Jollife (1996) The ecology of the avian brain: Food-storing memory and the hippocampus. Ibis, 138: 34–46.
40.
Lapointe, F.-J., G. Baron, and P. Legendre (1999) Encephalization, adaptation and evolution of chiroptera: A statistical analysis with further evidence for bat monophyly. Brain Behav. Evol., 54: 119–126.
41.
Legendre, P., and F.-J. Lapointe (1995) Matching behavioral evolution to brain morphology. Brain Behav. Evol., 45: 110–121.
42.
Mack, A. L. (1993) The sizes of vertebrate-dispersed fruits: a neotropical-paleotropical comparison. Am. Nat., 142: 840–856.
43.
Novacek, M.J. (1985) Evidence for echolocation in the oldest known bats. Nature, 315: 140–141.
44.
Pagel, M.D. (1992) A method for the analysis of comparative data. J. Theor. Biol., 156: 431–442.
45.
Pettigrew, J.D., B.G.M. Jamieson, S.K. Robson, L.S. Hall, K.I. McAnally, and H.M. Cooper (1989) Phylogenetic relations between microbats, megabats and primates (Mammalia: Chiroptera and Primates). Philos. Trans. R. Soc. Lond. B, 325: 489–559
46.
Robbins, L.W., and V.M. Sarich (1988) Evolutionary relationships in the family Emballonuridae (Chiroptera). J. Mamm., 69: 1–13.
47.
Simmons, N.B., and J.B. Geisler (1998) Phylogenetic relationships of Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx to extant bat lineages, with comments on the evolution of echolocation and foraging strategies in Microchiroptera. Bull. Am. Mus. Nat. Hist., 235: 1–182.
48.
Speakman, J.R (1999) The evolution of flight and echolocation in pre-bats: An evaluation of the energetics of reach hunting. Acta Chiropterol., 1: 3–15.
49.
Springer, M.S., E.C. Teeling, O. Madsen, M.J. Stanhope, and W.W. de Jong (2001) Integrated fossil and molecular data reconstruct bat echolocation. Proc. Natl. Acad. Sci., 98: 6241–6246.
50.
Szekely, A.D. (1999) The avian hippocampal formation: subdivisions and connectivity. Behav. Brain Res., 98: 219–225.
51.
Szekely, T., C.K. Catchppole, A. DeVoogd, Z. Marchl, and T.J. DeVoogd (1996) Evolutionary changes in a song control area of the brain (HVC) are associated with evolutionary changes in song repertoire among European warblers (Sylvidae). Proc. R. Soc. Lond. B, 263:607–610.
52.
Teeling, E.C., M. Scally, D.J. Kao, M.L. Romagnoli, M.S. Springer, and M.J. Stanhope (2000) Molecular evidence regarding the origin of echolocation and flight in bats. Nature, 403:188–192.
53.
Thies, W., E.K.V. Kalko, and H.-U. Schnitzler (1998) The roles of echolocation and olfaction in two Neotropical fruit-eating bats, Carollia perspicillata and C. castenea, feeding on Piper. Behav. Ecol. Sociobiol., 42: 397–409.
54.
Vanhooydonck, B., and R. Van Damme (1999) Evolutionary relationships between body shape and habitat use in lacertid lizards. Evol. Ecol. Res., 1: 785–805.
55.
von Helverson, D., and O. von Helverson (1999) Acoustic guide in bat-pollinated flower. Nature, 398: 759–760.
56.
von Helversen, O., L Winkler, and H.J. Bestmann (2000) Sulphur-containing ‘perfumes’ attract flower-visiting bats. J. Comp. Physiol. A., 186:143–153.
57.
Wetterer, A.L., M.V. Rockman, and N.B. Simmons (2000) Phylogeny of phyllostomid bats (Mammalia: Chiroptera): Data from diverse morphological systems, sex chromosomes, and restriction sites. Bull. Am. Mus. Nat. Hist., 248: 1–200.
Copyright / Drug Dosage / Disclaimer
Copyright: All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, microcopying, or by any information storage and retrieval system, without permission in writing from the publisher.
Drug Dosage: The authors and the publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accord with current recommendations and practice at the time of publication. However, in view of ongoing research, changes in government regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any changes in indications and dosage and for added warnings and precautions. This is particularly important when the recommended agent is a new and/or infrequently employed drug.
Disclaimer: The statements, opinions and data contained in this publication are solely those of the individual authors and contributors and not of the publishers and the editor(s). The appearance of advertisements or/and product references in the publication is not a warranty, endorsement, or approval of the products or services advertised or of their effectiveness, quality or safety. The publisher and the editor(s) disclaim responsibility for any injury to persons or property resulting from any ideas, methods, instructions or products referred to in the content or advertisements.
You do not currently have access to this content.