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	Author	Reader, S.M.; Laland, K.N.
	Title	Social intelligence, innovation, and enhanced brain size in primates			Type	Journal Article
	Year	2002	Publication	Proceedings of the National Academy of Sciences of the United States of America	Abbreviated Journal	Proc. Natl. Acad. Sci. U.S.A.
	Volume	99	Issue	7	Pages	4436-4441
	Keywords	Animals; Brain/anatomy & histology; Evolution; Intelligence; Learning; Primates/anatomy & histology/psychology; Social Behavior
	Abstract	Despite considerable current interest in the evolution of intelligence, the intuitively appealing notion that brain volume and “intelligence” are linked remains untested. Here, we use ecologically relevant measures of cognitive ability, the reported incidence of behavioral innovation, social learning, and tool use, to show that brain size and cognitive capacity are indeed correlated. A comparative analysis of 533 instances of innovation, 445 observations of social learning, and 607 episodes of tool use established that social learning, innovation, and tool use frequencies are positively correlated with species' relative and absolute “executive” brain volumes, after controlling for phylogeny and research effort. Moreover, innovation and social learning frequencies covary across species, in conflict with the view that there is an evolutionary tradeoff between reliance on individual experience and social cues. These findings provide an empirical link between behavioral innovation, social learning capacities, and brain size in mammals. The ability to learn from others, invent new behaviors, and use tools may have played pivotal roles in primate brain evolution.
	Address	Department of Zoology, University of Cambridge, High Street, Madingley, Cambridge CB3 8AA, United Kingdom
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0027-8424	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:11891325			Approved	no
	Call Number				Serial	2149
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	Author	Webster, M.M.; Laland, K.N.
	Title	Social learning strategies and predation risk: minnows copy only when using private information would be costly			Type	Journal Article
	Year	2008	Publication	Proc Biol Sci	Abbreviated Journal	Proc. R. Soc. Lond. B
	Volume	275	Issue	1653	Pages	2869-2876
	Keywords
	Abstract	Animals can acquire information from the environment privately, by sampling it directly, or socially, through learning from others. Generally, private information is more accurate, but expensive to acquire, while social information is cheaper but less reliable. Accordingly, the 'costly information hypothesis' predicts that individuals will use private information when the costs associated with doing so are low, but that they should increasingly use social information as the costs of using private information rise. While consistent with considerable data, this theory has yet to be directly tested in a satisfactory manner. We tested this hypothesis by giving minnows (Phoxinus phoxinus) a choice between socially demonstrated and non-demonstrated prey patches under conditions of low, indirect and high simulated predation risk. Subjects had no experience (experiment 1) or prior private information that conflicted with the social information provided by the demonstrators (experiment 2). In both experiments, subjects spent more time in the demonstrated patch than in the non-demonstrated patch, and in experiment 1 made fewer switches between patches, when risk was high compared with when it was low. These findings are consistent with the predictions of the costly information hypothesis, and imply that minnows adopt a 'copy-when-asocial-learning-is-costly' learning strategy.
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	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6196
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	Author	Boogert, N.J.; Reader, S.M.; Hoppitt, W.; Laland, K.N.
	Title	The origin and spread of innovations in starlings			Type	Journal Article
	Year	2008	Publication	Animal Behaviour	Abbreviated Journal	Anim. Behav.
	Volume	75	Issue	4	Pages	1509-1518
	Keywords	diffusion dynamics; dominance; foraging; group; innovation; neophobia; social learning; social network; starling; Sturnus vulgaris
	Abstract	There are numerous reports of novel learned behaviour patterns in animal populations, yet the factors influencing the invention and spread of these innovations remain poorly understood. Here we investigated to what extent the pattern of spread of innovations in captive groups of starlings, Sturnus vulgaris, could be predicted by knowledge of individual and social group variables, including association patterns, social rank orders, measures of neophobia and asocial learning performance. We presented small groups of starlings with a series of novel extractive foraging tasks and recorded the latency for each bird to contact and solve each task, as well as the orders of contacting and solving. We then explored which variables best predicted the observed diffusion patterns. Object neophobia and social rank measures characterized who was the first of the group to contact the novel foraging tasks, and the subsequent spread of contacting tasks was associated with latency to feed in a novel environment. Asocial learning performance, measured in isolation, predicted who was the first solver of the novel foraging tasks in each group. Association patterns did not predict the spread of solving. Contact latency and solving duration were negatively correlated, consistent with social learning underlying the spread of solving. Our findings indicate that we can improve our understanding of the diffusion dynamics of innovations in animal groups by investigating group-dependent and individual variables in combination. We introduce novel methods for exploring predictors of the origin and spread of behavioural innovations that could be widely applied.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6036
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	Author	Boogert, N.J.; Reader, S.M.; Laland, K.N.
	Title	The relation between social rank, neophobia and individual learning in starlings			Type	Journal Article
	Year	2006	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	72	Issue	6	Pages	1229-1239
	Keywords
	Abstract	Researchers with diverse interests in topics ranging from the formation of dominance hierarchies and social intelligence to animal personalities have predicted specific, and often conflicting, relations between social rank, neophobia and learning ability. We investigated the relations between these variables in captive groups of wild-caught starlings, Sturnus vulgaris, adopting a multidimensional approach to social rank and neophobia. Both agonistic and competitive rank orders were determined for each group and we tested individuals in the absence of their groupmates for object neophobia, latency to feed in a novel environment and performance on an extractive foraging task. In each starling group, the fastest learners occupied the highest competitive ranks, supporting the hypothesis that cognitive ability is positively correlated with social dominance. Competitive rank orders, however, did not correlate significantly with agonistic rank orders. Situation-specific foraging neophobia was suggested: individuals showed consistency in their latencies to feed near a variety of novel objects, but no significant correlation was found between this measure of object neophobia and latency to feed in a novel environment. Starlings fastest to feed in the novel environment were fastest in solving the foraging task. We discuss the implications of these findings for researchers studying hierarchy formation in animal groups, social intelligence and animal personalities.
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	Notes				Approved	yes
	Call Number	Equine Behaviour @ team @ room B 3.029			Serial	2074
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	Author	Reader, S.M.; Kendal, J.R.; Laland, K.N.
	Title	Social learning of foraging sites and escape routes in wild Trinidadian guppies			Type	Journal Article
	Year	2003	Publication	Animal Behaviour	Abbreviated Journal	Anim. Behav.
	Volume	66	Issue	4	Pages	729-739
	Keywords
	Abstract	We describe two field experiments with wild guppies, Poecilia reticulata, in Trinidad that demonstrated that guppies can acquire foraging and predator escape-response information from conspecifics. In the foraging experiment, subjects were presented with two distinctly marked feeders in their home rivers. One feeder contained a conspecific shoal in a transparent container. Guppies preferred to enter the feeder containing this artificial shoal over the other feeder. In a test phase, the artificial shoal was removed and the feeders replaced at the testing site after a 5-min delay. More guppies entered the feeder that had contained the artificial shoal over the other feeder, a difference that can be explained only by the fish learning the characteristics or location of the feeder during the training phase. We suggest that subjects acquired a foraging patch preference through a propensity to approach feeding conspecifics, a local enhancement process. In the predator escape-response experiment, naive 'observer' guppies could avoid an approaching trawl net by escaping through either a hole to which 'demonstrator' guppies had been trained or through an alternative hole. When the demonstrators were present, the naive observers escaped more often and more rapidly by the demonstrated route than the alternative route. When the demonstrators were removed, observers maintained a route preference according to the training of their demonstrators, which suggests that the observers had learned an escape route through following or observing their more knowledgeable conspecifics. Thus, both experiments reveal that guppies can socially learn in the wild. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6163
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	Author	Day, R.L.; Coe, R.L.; Kendal, J.R.; Laland, K.N.
	Title	Neophilia, innovation and social learning: a study of intergeneric differences in callitrichid monkeys			Type	Journal Article
	Year	2003	Publication	Animal Behaviour	Abbreviated Journal	Anim. Behav.
	Volume	65	Issue	3	Pages	559-571
	Keywords
	Abstract	In a comparative study of neophilia, innovation and social attentiveness we exposed individuals in seven callitrichid species, from three genera, to novel extractive foraging tasks. The results revealed consistently shorter response latencies, higher levels of successful and unsuccessful manipulation, and greater attentiveness to the task and to conspecifics inLeontopithecus (lion tamarins) than in both Saguinus (tamarins) and Callithrix (marmosets). This is consistent with the hypothesis that species dependent upon manipulative and explorative foraging tend to be less neophobic and more innovative than other species. Furthermore, Callithrix appeared to be less neophobic than Saguinus; ifCallithrix is regarded as the greater specialist, this result is inconsistent with the hypothesis that neophobia is associated with foraging specialization. We consider the relevance of our findings to taxonomic relationships, and to technical and Machiavellian intelligence hypotheses and discuss the implications for captive breeding and reintroduction strategies.Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.
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	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6035
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	Author	Galef, B.G.; Laland, K.N.
	Title	Social Learning in Animals: Empirical Studies and Theoretical Models			Type	Journal Article
	Year	2005	Publication	BioScience	Abbreviated Journal
	Volume	55	Issue	6	Pages	489-499
	Keywords
	Abstract	AbstractThe last two decades have seen a virtual explosion in empirical research on the role of social interactions in the development of animals' behavioral repertoires, and a similar increase in attention to formal models of social learning. Here we first review recent empirical evidence of social influences on food choice, tool use, patterns of movement, predator avoidance, mate choice, and courtship, and then consider formal models of when animals choose to copy behavior, and which other animals' behavior they copy, together with empirical tests of predictions from those models.
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	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
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	Series Volume		Series Issue		Edition
	ISSN	0006-3568	ISBN		Medium
	Area		Expedition		Conference
	Notes	10.1641/0006-3568(2005)055[0489:Sliaes]2.0.Co;2			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6398
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	Author	Laland, K.N.; Reader, S.M.
	Title	Foraging innovation in the guppy			Type	Journal Article
	Year	1999	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	57	Issue	2	Pages	331-340
	Keywords
	Abstract	When novel behaviour patterns spread through animal populations, typically one animal will initiate the diffusion. It is not known whether such 'innovators' are particularly creative individuals, individuals exposed to the appropriate environmental contingencies, or individuals in a particular motivational state. We describe three experiments that investigated the factors influencing foraging innovation in the guppy Poecilia reticulata. We exposed small laboratory populations of fish to novel foraging tasks, which involved exploration and problem solving to locate a novel food source. Experiments 1 and 2 found that (1) females were more likely to innovate than males, (2) food-deprived fish were more likely to innovate than nonfood-deprived subjects, and (3) smaller fish were more likely to innovate than larger fish. We suggest that the sex difference may reflect parental investment asymmetries in males and females. Experiment 3 found that past innovators were more likely to innovate than past noninnovators. Collectively, the results suggest that differences in foraging innovation in guppies are best accounted for by differences in motivational state, but, in addition, guppies may vary in their predisposition to innovate. Copyright 1999 The Association for the Study of Animal Behaviour.
	Address	Sub-Department of Animal Behaviour, Department of Zoology, University of Cambridge
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:10049472			Approved	no
	Call Number				Serial	2151
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	Author	Reader, S.M.; Laland, K.N.
	Title	Diffusion of foraging innovations in the guppy			Type	Journal Article
	Year	2000	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	60	Issue	2	Pages	175-180
	Keywords
	Abstract	The way in which novel learned behaviour patterns spread through animal populations remains poorly understood, despite extensive field research and the recognition that such processes play an important role in the behavioural development, social interactions and evolution of many animal species. We conducted a series of controlled diffusions of foraging information in replicate experimental populations of the guppy, Poecilia reticulata. We presented novel foraging tasks over 15 trials to mixed-sex groups, made up of food-deprived and nonfood-deprived adults (experiment 1) or small, young fish and old, large adults (experiment 2). In these diffusions, knowledge of a route to a feeder could spread through the group by subjects learning from others, discovering the route for themselves, or, most likely, by some combination of these social and asocial learning processes. We found a striking sex difference, with novel foraging information spreading at a significantly faster rate through subgroups of females than of males. Females both discovered the goal and learned the route more quickly than males. Food-deprived individuals were faster at completing the tasks over the 15 trials than nonfood-deprived guppies, and there was a significant interaction between sex and size, with a sex difference in adults but not young individuals. There was also an interaction between sex and hunger level, with food deprivation having a stronger effect on male than female performance. We suggest that information may diffuse in a similar nonrandom or 'directed' manner through many natural populations of animals. Copyright 2000 The Association for the Study of Animal Behaviour.
	Address	Sub-Department of Animal Behaviour, Department of Zoology, University of Cambridge
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:10973718			Approved	no
	Call Number				Serial	2150
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	Author	Kendal, R.L.; Coe, R.L.; Laland, K.N.
	Title	Age differences in neophilia, exploration, and innovation in family groups of callitrichid monkeys			Type	Journal Article
	Year	2005	Publication	American journal of primatology	Abbreviated Journal	Am. J. Primatol.
	Volume	66	Issue	2	Pages	167-188
	Keywords	Age Factors; Analysis of Variance; Animals; Animals, Zoo; Behavior, Animal/physiology; Callitrichinae/physiology; Creativeness; Exploratory Behavior/*physiology; Observation; Social Behavior; Task Performance and Analysis
	Abstract	The prevailing assumption in the primate literature is that young or juvenile primates are more innovative than adult individuals. This innovative tendency among the young is frequently thought to be a consequence, or side effect, of their increased rates of exploration and play. Conversely, Reader and Laland's [International Journal of Primatology 22:787-806, 2001] review of the primate innovation literature noted a greater reported incidence of innovation in adults than nonadults, which they interpreted as (in part) a reflection of the greater experience and competence of older individuals. Within callitrichids there is contradictory evidence for age differences in response to novel objects, foods, and foraging tasks. By presenting novel extractive foraging tasks to family groups of callitrichid monkeys in zoos, we examined, in a large sample, whether there are positive or negative relationships of age with neophilia, exploration, and innovation, and whether play or experience most facilitates innovation. The results indicate that exploration and innovation (but not neophilia) are positively correlated with age, perhaps reflecting adults' greater manipulative competence. To the extent that there was evidence for play in younger individuals, it did not appear to contribute to innovation. The implications of these findings for the fields of innovation and conservation through reintroduction are considered.
	Address	Sub-Department of Animal Behavior, University of Cambridge, Cambridge, UK. RachelKendal2003@yahoo.co.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0275-2565	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15940712			Approved	no
	Call Number				Serial	2148
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