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J. U . M. (1978). Energetics of survival in Hererocephahs &her (Ruppell). the naked molerat (Rodentia: Bathyergidae). Bull. CarneRie Mus. Nar. Hist. 6 , 81-87. Jarvis, J. U. M. (1981). Eusociality in a mammal: Cooperative breeding in naked mole-rat colonies. Science 212, 571-573. Jarvis. J. U. M . , and Sale, J. B. (1971). Burrowing and burrow patterns of East African mole-rats Tachvorvcres. Heliophobius and Hererocephalus. J . Zool. 163, 45 1-479. Jeanne, R. L. (1979). Construction and utilization of multiple combs in Polisres canadensis in relation to the biology of a predaceous moth.
Similar effects of helpers have been described in mammals (Moehlman, 1979). There were several ambiguities in the interpretation of these studies. These were finally all controlled in an experimental study of the effects of removal of helpers (Brown and Brown, 1981b; Brown et a l . , 1982), which reinforced the general finding that helpers do measurably increase reproductive success, at least in a species for which helping seems to be regular, frequent, and widespread (level c). 4. What is the Relative Importance of D, d, I, and i ?
Rev. Ecol. Sysr. 3, 193-232. Hamilton, W. D. (1975). Innate social aptitudes of man: An approach from evolutionary genetics. I n “Biosocial Anthropology” (R. ), pp. 133-155. Wiley, New York. Hardy. J. W. (1976). Comparative breeding behavior and ecology of the bushy-crested and Nelson san bkds jay. Wilson Bull. 88, ii, 96-120. Hegner, R. , Emlen, S. T . , and Demong, N. J. (1982). Spatial organization of the white-fronted bee-eater Nature (London) 298, 264-266. , and Michener, C. D. (1980). Mechanisms of identification and discrimination in social hymenoptera.