On the number of subjects used in animal behaviour experiments

Experiments published in Animal Behaviour in 1979 are reviewed, and it is concluded that there are grounds for thinking that the numbers of animals used could sometimes be reduced without loss of scientific rigour. Principles underlying the choice of number of subjects (N) are outlined, and illustrated with examples from the 1979 sample. Several alternatives to increasing N in order to increase power are given, including better design, larger α, and sequential methods. The logic of inference from experiments with small N is discussed.     

Effect of resource competition on the long-term allocation of grooming by female baboons: evaluating Seyfarth's model

Seyfarth (1977, Journal of Theoretical Biology, 65, 671-698) proposed an influential model that explained the long-term patterning of grooming relationships between female primates in terms of an interaction between the idealized grooming objectives of females and competition for valuable grooming partners. A critical test of the model requires a demonstration not only that competition for partners exists, but also that females do have an underlying target for the amount of grooming that they would like to receive relative to the amount they need to give. As it is not possible to stipulate a priori what this target is, or to detect it directly, we assessed its applicability by making a set of predictions, within the framework of Seyfarth's model, as to how observed grooming patterns should change with changes in the intensity of resource competition. We tested these predictions with data from a troop of baboons, Papio hamadryas ursinus, for which the within-bout structure of grooming has already been shown to be sensitive to changes in resource competition (Barrett et al. 2002, Animal Behaviour, 63, 1047-1053). We found no evidence of competition for grooming partners and our results do not support the predictions of the modified model with respect to the existence of underlying grooming objectives. The findings that the grooming of female baboons is more diverse and that the mean rank distance separating partners increases when resource competition is greater, together with the absence of rank effects on long-term patterns, are, however, supportive of the recent biological market approach to social interactions. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Long- and short-term state-dependent foraging under predation risk: an indication of habitat quality

Animals living in environments of different quality will have different expectations of their future reproductive success and survival. This may affect the individual's risk-taking behaviour as manifest in the cost of predation. We investigated the foraging behaviour of starlings, Sturnus vulgaris, when perceived predation risk varied between patches. Short-term food availability varied between treatments and long-term differences in perceptions of environmental quality varied between groups of individuals. This corresponds to variation in the three components of the cost of predation (P): the predation risk (μ); the change in reproductive value with energy gain (∂F /∂e); and the reproductive value or fitness factor (F). The birds showed that they experienced a higher cost of predation while using the risky food patches (μ component) and in the high food treatment (∂F /∂e component). Furthermore, birds from a high-reward habitat revealed a higher P than birds from a poor habitat (F component). The results show that the costs of predation are possible to tease apart by using behavioural indicators. The method presented allows measurement of fitness prospects of individuals, which may have consequences for conservation, for example, to identify low-quality habitat. Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

The making of ethology: The association for the Study of Animal Behaviour, 1936–1986

The paper reviews the history of the Association for the Study of Animal Behaviour in its 50th anniversary year. Having discussed the founding of the Institute for the Study of Animal Behaviour in 1936, the paper establishes its principal aims, follows its fortunes before and during the war, traces its role in the expansion and institutionalization of ethology in Britainbetween 1945 and 1950, and summarizes its transformation into a flourishing scientific society in the 1950s and 1960s. It is concluded that the history of the Association is, to a striking extent, a microcosm of the making of ethology as a successful scientific discipline.     

Female-offspring communication in a Taiwanese tree frog, Chirixalus eiffingeri (Anura: Rhacophoridae)

We assessed the roles of visual and olfactory cues in female-tadpole communication inChirixalus eiffingeri . The mean cumulative time that at least one tadpole was active or begged for food was significantly longer when a female C. eiffingeri was present than when a plastic frog was introduced and when no frog was present. Tadpoles did not respond visually to a female frog physically separated from them by transparent Plexiglas. However, tadpoles were more active in water conditioned by female frogs than in unconditioned water. Tadpole activity was further elevated by water conditioned by a female frog and tadpoles. Tadpoles were more active in water conditioned by male frogs than in unconditioned water, but water conditioned by a male frog and tadpoles did not further elevate tadpole activity. Thus, water conditioned by adults of either sex contains substances that increase tadpole activity, but only females show a synergistic effect with conditioning by tadpoles. Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Taxonomic chauvinism revisited: insight from parental care research

Parental care (any non-genetic contribution by a parent that appears likely to increase the fitness of its offspring) is a widespread trait exhibited by a broad range of animal taxa. In addition to influencing the fitness of parent(s) and offspring, parental care may be inextricably involved in other evolutionary processes, such as sexual selection and the evolution of endothermy. Yet, recent work has demonstrated that bias related to taxonomy is prevalent across many biological disciplines, and research in parental care may be similarly burdened. Thus, I used parental care articles published in six leading journals of fundamental behavioral sciences (Animal Behaviour, Behavioral Ecology, Behavioral Ecology and Sociobiology, Ethology, Hormones and Behavior, and Physiology & Behavior) from 2001–2010 (n = 712) to examine the year-to-year dynamics of two types of bias related to taxonomy across animals: (1) taxonomic bias, which exists when research output is not proportional to the frequency of organisms in nature, and (2) taxonomic citation bias, which is a proxy for the breadth of a given article—specifically, the proportion of articles cited that refer solely to the studied taxon. I demonstrate that research on birds likely represents a disproportionate amount of parental care research and, thus, exhibits taxonomic bias. Parental care research on birds and mammals also refers to a relatively narrow range of taxonomic groups when discussing its context and, thus, exhibits taxonomic citation bias. Further, the levels of taxonomic bias and taxonomic citation bias have not declined over the past decade despite cautionary messages about similar bias in related disciplines— in fact, taxonomic bias may have increased. As in Bonnet et al. (2002), my results should not be interpreted as evidence of an ‘ornithological Mafia’ conspiring to suppress other taxonomic groups. Rather, I generate several rational hypotheses to determine why bias persists and to guide future work.     

Misprescription and misuse of one‐tailed tests

One‐tailed statistical tests are often used in ecology, animal behaviour and in most other fields in the biological and social sciences. Here we review the frequency of their use in the 1989 and 2005 volumes of two journals (Animal Behaviour and Oecologia), their advantages and disadvantages, the extensive erroneous advice on them in both older and modern statistics texts and their utility in certain narrow areas of applied research. Of those articles with data sets susceptible to one‐tailed tests, at least 24% in Animal Behaviour and at least 13% in Oecologia used one‐tailed tests at least once. They were used 35% more frequently with nonparametric methods than with parametric ones and about twice as often in 1989 as in 2005. Debate in the psychological literature of the 1950s established the logical criterion that one‐tailed tests should be restricted to situations where there is interest only in results in one direction. ‘Interest’ should be defined; however, in terms of collective or societal interest and not by the individual investigator. By this ‘collective interest’ criterion, all uses of one‐tailed tests in the journals surveyed seem invalid. In his book Nonparametric Statistics, S. Siegel unrelentingly suggested the use of one‐tailed tests whenever the investigator predicts the direction of a result. That work has been a major proximate source of confusion on this issue, but so are most recent statistics textbooks. The utility of one‐tailed tests in research aimed at obtaining regulatory approval of new drugs and new pesticides is briefly described, to exemplify the narrow range of research situations where such tests can be appropriate. These situations are characterized by null hypotheses stating that the difference or effect size does not exceed, or is at least as great as, some ‘amount of practical interest’. One‐tailed tests rarely should be used for basic or applied research in ecology, animal behaviour or any other science.     

Nest-building activity and laying date influence female reproductive investment in magpies: an experimental study

Nest size or nest-building activity has recently been hypothesized to be a postmating sexually selected signal in monogamous birds: females may assess a male's parental quality and willingness to invest in reproduction by his participation in nest building. Females may thus adjust their reproductive effort (i.e. clutch size) not only to their own abilities but also to those of their mates. We investigated whether female magpies, Pica pica, use nest-building activity rather than nest size to adjust their reproductive effort during replacement breeding attempts. After we removed their first clutch, high-quality pairs that built a large nest for the first clutch were more capable of building a replacement nest and females adjusted their clutch size in relation to the time it took to build the nest rather than nest size. We also found support for the hypothesized trade-off between clutch size and egg size in magpies. In replacement clutches females decreased clutch size and increased egg volume, thereby probably improving the survival probability of their offspring in less favourable conditions.Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Context-dependent visual preferences in starlings and blue tits: mate choice and light environment

There is considerable interest in the role that ultraviolet (UV) cues play in the foraging and mate choice decisions of birds. However, with the exception of the zebra finch, Taeniopygia guttata, it is not yet clear whether ultraviolet preferences are context specific, or whether birds show a general preference for full-avian-spectrum environments (320-700 nm) irrespective of the activity in which they are engaged. We investigated whether European starlings, Sturnus vulgaris, and blue tits, Parus caeruleus, show general (nonresource based) or context-specific preferences for full-spectrum environments. We found that neither species showed a general preference for UV-present (UV+) over UV-deficient (UV−) environments, when those environments contained no resources (experiment 1). Furthermore, neither species showed a UV+ preference when cages contained food, water and perches (starlings; experiment 2) or food, perches and heterospecifics (blue tits; Hunt et al. 1999. Animal Behaviour, 58, 809-815). However, both species did show highly significant preferences for UV+ conditions when viewing potential mates. Such experiments are necessary before one can conclude that particular wavebands have specific relevance to mate choice. In fact, our results suggest that the importance of particular wavelength compositions do indeed vary with behavioural context.     

Adaptation to a novel host modifies host discrimination by the seed beetle Callosobruchus maculatus

Females of the seed beetle Callosobruchus maculatus avoid adding eggs to seeds that already bear eggs. Geographical variation in this behaviour has been thought to depend on differences in host size. In populations that attack small-seeded legumes, only one or two larvae can develop within a seed, and females are especially adept at detecting and rejecting occupied (egg-laden) seeds. We performed a mass-selection experiment in which replicate lines of a population associated with a small host (mung bean, Vigna radiata) were either maintained on this host or were transferred to a larger host (cowpea, Vigna unguiculata) that can support several larvae per seed. After more than 40 generations, we estimated the strength of host discrimination by presenting females a choice of egg-free and egg-laden seeds, and by quantifying how uniformly females spread their eggs among egg-free seeds. Compared to females maintained on mung bean, females from cowpea lines were more likely to accept occupied seeds in choice tests. They also distributed their eggs less uniformly, especially when cowpea served as the test host. Cowpea lines thus evolved to resemble populations that have long been associated with the larger host. A separate study showed that weaker host discrimination in the cowpea lines was accompanied by a decline in larval competitiveness, which may have further relaxed selection for avoidance of occupied hosts. Our results demonstrate that switching to a novel resource can produce rapid and predictable changes in a fitness-related insect behaviour. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Queen reproduction, chemical signalling and worker behaviour in polygyne colonies of the ant Formica fusca

Potential reproductive conflicts are common in social insects and may occur between and within castes. In multiqueen (polygyne) colonies of ants, reproductive conflicts among coexisting queens may be resolved by aggressive interactions and/or by pheromonal signalling. Pheromonal signals may be directed at workers, which may adjust the reproductive shares of queens behaviourally. Workers are expected to favour a queen that is more likely to be their mother and to produce offspring of close kin to them. We used microsatellite markers to study reproductive partitioning between cobreeding Formica fusca queens in a laboratory experiment where workers received a choice between two queens. We expected queens of this species to communicate their reproductive status by chemical communication, because no aggressive interactions between queens have been observed. We found that queens of different reproductive status (with majority, minority or no production) differed in their cuticular hydrocarbon (CHC) profiles. The fecundity of a queen was associated with worker behaviour; the higher a queen's fecundity, the more attention she received from workers. Our results suggest that a queen fecundity signal is encoded in her CHC profile and acts as a pheromone for workers, which respond to the signal by discriminating between queens. Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Muting individual nestlings reduces parental foraging for the brood

Nestling birds use vocal and visual behaviours when soliciting food from parents. Such behaviours serve at least two discrete functions: (1) to induce parents to bring more food; and (2) to influence how food is allocated among brood members. Playback experiments have shown that vocalizations serve function 1. Do they also function to influence intrabrood allocation, as contemporary begging theory suggests, or is that governed chiefly by the nonvocal components of begging (neck stretching, gaping, jockeying for position within the nest)? We tested this hypothesis using a novel nonsurgical muting technique to decouple the vocal and visual components of begging in nestling red-winged blackbirds, Agelaius phoeniceus. Single chicks that were muted temporarily (1 h) continued to be fed at roughly the same rate as either the same individual prior to muting or sham-muted nestlings in the same brood. Parents reduced provisioning rates by increasing nest attentiveness in response to changes in the begging behaviour of the brood following treatment. These changes included less time spent begging (visual and vocal) accompanied by a reduction in the collective vocalizations of the brood. Our results suggest that vocalizations function primarily to regulate parental foraging rates, and visual begging displays function primarily to access food (competition).Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Is a cultural ethology possible?

The possibility, desirability, and potential outcomes of applying ethological methods to the study of culture-specific human behaviors are investigated. Ethology and culture are explored. A new term, instruction, and its use in cultural ethology are proposed. Genetics and survival value are related to cultural ethology. A cultural ethology is given a possible theoretical foundation, and current attempts at a cultural ethology are appraised. A research program in cultural ethology and related fields is proposed.This article is a completely rewritten version of a paper of the same title delivered at the annual meeting of the Southern Anthropological Society at Gainesville, Florida, in February 1968 and published inResearch Previews 15(1): 37–47, April 1968.     

Material and genetic benefits of female multiple mating and polyandry

The maintenance of female polyandry has traditionally been attributed to the material (direct) benefits derived from male mating resources (e.g. nuptial gifts) accrued by multiple mating. However, genetic (indirect) benefits offer a more robust explanation since only polyandrous, not monandrous, females may gain both material benefits from multiple mating and genetic benefits from multiple sires. Discriminating between material and genetic benefits is essential when addressing the mechanism by which polyandry is adaptively maintained, but are difficult to disentangle because they affect fitness in similar ways. To test the hypothesis that genetic benefits maintain polyandry, we compared four components of fitness (longevity, fecundity, hatching success and survivorship) between monandrous and polyandrous females in the ground cricket, Allonemobius socius. We discovered that females derived nongenetic benefits from mating multiply, in that the magnitude of the nuptial gift was positively associated with the number of eggs produced. However, polyandrous females had over a two-fold greater hatching success and a 43% greater offspring survivorship, leading to a significantly higher relative fitness than the monandrous strategy. These results were independent of the confounding effects of material benefits, implying that genetic contributions play a large role in the maintenance of polyandry and potentially in the antagonistic coevolutionary relationship between polyandry and male nuptial gifts. Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Maternal care in the rockpool amphipod Apherusa jurinei: developmental and environmental cues

There is evidence that active, pre-emergence maternal brood care in amphipod crustaceans may be associated with ‘harsh’ environmental conditions. We examined, in the rockpool amphipod Apherusa jurinei, behavioural activities that may function as a form of active brood care. Only ovigerous females showed ‘curl’ and ‘stretch’ activities, with consequent flushing of the brood pouch and cycling of the eggs therein. There was a significant decline in these activities as embryonic development advanced and brood care almost ceased when well-developed embryos showed a heart pulse and self-ventilation. We propose that this pattern of brood care reflects changes in the physiological requirements of embryos as they develop within the egg membrane. In addition, ovigerous females showed significantly higher levels of brood care under lowered oxygen conditions. They achieved this by increasing the average duration of the ‘stretch’ component, with other brood care components remaining constant. Thus, developmental and environmental cues alter the components of active brood care in distinct ways. Experimental removal showed that the physical presence of eggs in the brood pouch is important in controlling the expression of brood care activities. However, females with all of their eggs removed continued to brood at low levels, suggesting that a maternal state also controls brood care. The sophisticated expression of active maternal brood care in amphipods under ‘harsh’ environmental conditions such as rockpools has implications both for individual reproductive success and the distribution and abundance of brooding versus nonbrooding species. Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

The Running,Jumping and Standing-still Game

Films Hydra: Feeding Behaviour BGP781 Hydra: Reproductive Behaviour BGP782 Two 8 mm, colour, loop films, 3.5 minutes each, with guidance notes. Prices: Super 8 £13.00 each; Standard 8 £12.30 each. Available from Gateway Educational Media, Waverly Road, Yale, Bristol BS17 5RB; Mark Thomas

Slides Hydra TZ6 Set of 11, 35 mm slides by S. Walker (1969). Price £7.20. Ref. M33625/3. Available from Philip Harris Biological Ltd, Oldmixon, Weston-Super-Mare, Avon BS24 9BJ; Janet Croston

Slides Form and function in flowering plants B10/003 Fitness for survival B10/005 30-framefilmslrips with cassette tapes and brief notes; running times 28 minutes. Price £10.00 each. Available from Audio Learning Ltd, 183–189 Queensway, London W2 5HL; Ruth Dormandy, Jane Harding, Brian Kenway, Andrew Richards

Charts The Animal Kingdom The Plant Kingdom Wall charts, 66 cm by 50 cm, printed in full colour, by Pamela Haddon and Don Mackean. Price £1.60. ISBN 0 7195 3834 3. Published by John Murray Ltd, 50 Albemarle Street, London WIX4BD; John A. Barker

Charts Lichens and Air Pollution Chart, 80 cm by 60 cm, in full colour. UK Education price £1.00. Reference WI/1. Available from BP Educational Service, PO Box 5, Welherby, West YorksLS23 7EH; John A. Barker

Charts Befriend a friend A set of six black-and-white posters produced by the RSPCA Schools Campaign for Animal Welfare. Available from the RSPCA, Purchase and Supply Department, Causeway, Horsham, West Sussex RH12 1HG. Price 15p each; C. Dale     

Evolution of female colour polymorphism in damselflies: testing the hypotheses

The existence of several female colour morphs is a conspicuous characteristic of many damselflies that show one male-like (androchrome) and several nonmale-like (gynochrome) morphs. We tested several adaptive hypotheses and the null model for the maintenance of female polychromatism (one androchrome and two gynochromes) in the damselfly Ceriagrion tenellum. We tested the null model by comparing the degree of genetic differentiation between the colour locus and a set of 19 neutral RAPD loci in five populations. Our results indicate that selection is acting to maintain similar frequencies between populations at the colour locus. Using mark–recapture techniques we found that mating success is not dependent on female coloration. We tested the mimicry hypothesis by presenting live and dead models to males. Dead models were highly attractive irrespective of coloration. In contrast, with live models males could not distinguish between androchromes and other males, and were more attracted to gynochrome females. Despite this, within populations morph frequencies remained constant over time and mating was at random with respect to female coloration. However, there was a positive relationship between male density and androchrome frequency in a comparative study of eight populations. We discuss our results in the framework of sexual conflict theory and suggest that andro- and gynochrome females are using different strategies to control their number of matings. The different morphs might be maintained in a balanced polymorphism by a combination of density- and frequency-dependent mechanisms.Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

The Ibis: Transformations in a Twentieth Century British Natural History Journal

The contents of the British Ornithologists' Union's journal, The Ibis, during the first half of the 20th century illustrates some of the transformations that have taken place in the naturalist tradition. Although later generations of ornithologists described these changes as logical and progressive, their historical narratives had more to do with legitimizing the infiltration of the priorities of evolutionary theory, ecology, and ethology than analyzing the legacy of the naturalist tradition on its own terms. Despite ornithologists' claim that the journal's increasing focus on "biology" represented a natural development after the preliminary phase of systematics and geographical ornithology, in fact a small group campaigned to bring the priorities of population ecology, behavior, and selection theory into the journal and British ornithology more generally. The problems involved in this transition highlight the importance of methodological and institutional context in determining and reinforcing appropriate research programs for ornithologists. Comparing the discipline-building rhetoric of moderns with the contents of the past illustrates how modern evaluations of 19th century research programs have been enmeshed in ornithologists' endeavors to forge new identities for traditional disciplines.