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Cognition in Woodpeckers 63-76 05Gajdon Layout 2 30.11.15 15:49 Seite 63 ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Denisia Jahr/Year: 2015 Band/Volume: 0036 Autor(en)/Author(s): Gajdon Gyula K. Artikel/Article: Cognition in woodpeckers 63-76 05Gajdon_Layout 2 30.11.15 15:49 Seite 63 Cognition in woodpeckers Gyula K. G AJDON & Hans W INKLER Abstract : Despite their large telencephalons, not much is known about cognition in woodpeckers. The hypothesis that large brains evolved in the context of a complex social life seems not to hold for most woodpeckers because many large brained species are rather solitary. Nevertheless, woodpeckers show signs of flexible communicative skills. Recently, the social brain hypothesis has been challenged by broader concepts of the relevance of behavioral flexibility for the evolution of large brains. Especially in the context of extractive foraging, woodpecker may require a good spatial memory and sophisticated technical skills. Woodpeck - ers rank high among birds with respect to feeding innovations. The extractive foraging style of woodpeckers may require high motivation to explore what in turn could positively affect cognitive performance. Key words : Animal cognition, social brain hypothesis, extractive foraging, technical intelligence, sensorimotor intelligence, anvils, reversal learning, Picidae. Introduction The evolution of Birds provide especially interesting subjects for ideas on animal cognition studying cognition, the brain and their adaptive varia - Cognition, learning, tool use tion because large brains evolved independently more The topic of cognition, especially in birds, has a often in birds, and in fact other sauropsids, than in long and complicated history in the behavioral sciences mammals ( NEALEN & RICKLEFS 2001; ISLER & V AN (DELIUS et al. 2000). This will be outlined in this, and SCHAIK 2009; BALANOFF et al. 2013). This seems to be examples from the research on woodpeckers will illus - linked with an altricial breeding style that evolved sev - trate it further. Although it has been known for some eral times in birds and that helps to buffer the costs of time that woodpecker have relatively large brains, rela - developing a large brain ( BENNETT & H ARVEY 1985; tively little is known about their cognitive performance RICKLEFS & S TARCK 1998; IWANIUK & N ELSON 2003). when compared to corvids, parrots, and even the small Because brain and body size are correlated, most brained pigeons or chickens. Only recently the evolu - authors postulate that brain size has to be related to tionary ecologist Robert RICKLEFS seemed to have body size for comparing brains and other traits across become aware of this fact. When reviewing the evi - species. For instance, residuals obtained from a log-log dence on the association between certain aspects of life regression of brain size on body size provide a useful history and brain size in birds, he wrote “I must have measure of relative brain size (see WINKLER & WIN - underestimated woodpeckers in the past …” RICKLEFS KLER , this volume). Regression analyses indicate that (2004, p. 123). crows, ravens, and jays (Corvidae), parrots (Psittaci - dae), hornbills (Bucerotidae), owls (Strigidae), and the This complex history of cognition research is also true woodpeckers (Picinae) have larger than average partially grounded in the nomenclature of the avian brain because brain regions considered to process cog - brains ( MLÍKOVSKÝ 1989; see WINKLER & WINKLER , this nitively more demanding tasks in mammals were volume). For example the Great Spotted Woodpecker wrongly homologized (The Avian Brain Nomenclature has about 1.6 times more brain volume than the about Consortium 2005). Previously, it was believed that equally sized blackbird ( Turdus merula ) (see WINKLER & these areas are relatively small in birds compared to WINKLER , this volume). mammals. Cortical areas are not as easily identified in The purposes of this paper are to review our current avian brains as in mammals due to a different cyto- knowledge about the cognitive performance of wood - architecture in birds. The unique mammalian cortex Denisia 36 , peckers, to consider some of the factors selecting for consists of six layers and thus differs clearly from other zugleich Kataloge des oberösterreichischen high cognitive performance and brain size and, finally, brain areas; while in birds comparable structures Landesmuseums Neue Serie 164 (2015): present some results of a pilot study on reversal learning. exhibit only three layers if any ( JARVIS 2009). In 2005, 63-76 05Gajdon_Layout 2 30.11.15 15:49 Seite 64 a large consortium of authors published a revised researchers increasingly regard cognition as an adaptive nomenclature of the avian brain, rendering the bird response to rather specific ecological problems. Among telencephalon on par with that of mammals (The Avian other things, scientists tested the hypothesis that large Brain Nomenclature Consortium 2005). And recent brains evolved to find adaptive answers to rather novel reviews indicate not only some important differences, and unusual challenges; what Daniel SOL called the cog - but also surprising similarities in the neural circuits in nitive buffer hypothesis ( SOL et al. 2007, SOL 2009). birds and mammals and discuss the consequences for Interestingly, this shift of emphasis is linked with understanding their cognitive processes ( EMERY & discussions between those scientists interested in pri - CLAYTON 2004; BUTLER & COTTERILL 2006, BUTLER et mates and those in birds and their respective interpreta - al. 2005; DUGAS -F ORD et al. 2012). These recent tion of research results. For example, primates’ primacy anatomical and neurological findings contributed to a in tool using was challenged by the sophisticated way general change in attitude from a pejorative “bird brain” New Caledonian Crows ( Corvus moneduloides ), use and to “brainy birds”. modify tools in the wild ( ORENSTEIN 1972; HUNT 1996, Comparative neuroanatomy and the comparative 2003). One captive New Caledonian crow was found to study of behavior have long histories reaching back to spontaneously bend a piece of wire in order to form a DARWIN (1872). In particular animal learning is one of hook for solving a new problem ( WEIR et al. 2002). It the oldest topics studied in animal behavioral sciences seems that physical cognition in New Caledonian because learning was of major interest in the psycholog - Crows may have become adapted to tool-using behav - ical sciences right from their beginning ( MACKINTOSH ior. But later on it was found that captive Rooks ( Corvus 1974). Psychologists have traditionally used rats as the frugilegus ) that do not use tools commonly in the wild, main model species. Ethology, behavioral and evolu - solved the same task of bending a wire into a hook with tionary ecology, as well as behavioral biology, neuroe - a high level of consistency ( BIRD & E MERY 2009). This cology and affective neuroscience, became flourishing raised the yet unsolved question of whether tool-using disciplines later on, and accordingly the main emphasis behavior in New Caledonian Crows, rather than being about the nature of cognition changed as these disci - an adaptive cognitive specialization, is a result of gen - plines evolved. Thus, while at first animal cognition eral intelligence in large brained crows ( CNOTKA et al. largely referred to learning mechanisms investigated by 2008; KACELNIK 2009; LIND et al. 2009). psychologists ( THORPE 1956), its concepts widened and The Social Brain Hypothesis evolved. According to Sara SHETTLEWORTH , a behav - ioral ecologist, cognition refers to ways in which ani - The Social Brain Hypothesis (SBH) dominated the mals (including humans) retain, process, and act on field of animal cognition for some decades. It claims that intelligence evolved in response to social rather information taken in through the senses ( SHETTLE - than ecological problems in primates in particular WORTH 2001; DUKAS 2004). It includes processes such as perception, learning, memory, and problem solving. (DUNBAR 1998; ADOLPHS 1999; PÉREZ -B ARBERÍA et al. This broad definition refers to the interest in the mod - 2007). The rationale of the SBH is that social life in ularity of information processing. Anthropologist larger primate groups is especially demanding because social partners are complex agents in terms of how they Duane RUMBAUGH and colleagues provide a definition in the narrower sense by which cognition refers to will interact and how they can be exploited, alone or “knowing, the creative capacity to reorganize percep - together with allies, against other allied individuals ( DE WAAL 2007). Keeping track of what all dyads did, where tion and past learning to generate new solutions to and with whom, rapidly increases computational problems” ( RUMBAUGH et al. 1996) Thus, this proposal, demands as group size increases. Accordingly, there was despite being aware of the importance of perception, much interest in the scientific community to demon - restricts cognition to some forms of problem solving by strate and dissect mechanisms of social learning, that is the reorganization of mental representations of the the modification of behavior by witnessing what others environment in the tradition of the cognitive revolu - are doing ( WHITEN , et al. 2004). Imitation, the ability tion as a counter movement to the models of associative to copy the form of a complete action on the same learning that behaviorists had
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