Novelty Responses in a Bird Assemblage Inhabiting an Urban Area

Neophobia, or the hesitancy to approach a novel food item, object, or place, is an important factor influencing the foraging behavior of animals. Environmental factors (e.g. rapid anthropogenic changes, migration into new habitats) are associated with novelty in feeding ecology and may affect neophobic responses. Mechanisms that underlie the differential neophobic response may involve complex interactions with the environment: post-fledging experience in a greater diversity of habitats or in habitats that are more complex may contribute to reduced neophobia. In a previous study, it was observed that some urbanized species, in particular house sparrows ( Passer domesticus ) and shiny cowbirds ( Molothrus bonariensis ) show high levels of neophobia. This study was carried out in a suburban marsh of Cortaderia selloana , a relatively simple and predictable ecosystem as compared to urban areas. For this reason, in the present study, we explored novelty responses of bird species inhabiting an urban area, representing a complex environment. The results were compared to those obtained previously in the suburban marsh. We found unexpectedly high levels of neophobia in house sparrows, but shiny cowbirds showed a somewhat neophilic response. In the presence of novel objects, house sparrows tended to enter the feeders alone, while shiny cowbirds tended to forage in groups. We found no differences in latencies to forage or in visit duration between habitat types, but the proportion of individuals that visited the feeders when novel objects were present was lower in the urban area for house sparrows and eared doves ( Zenaida auriculata ). The results are discussed in the context of invasion success and feeding innovation in shiny cowbirds.     

Migratory New World Blackbirds (Icterids) Are More Neophobic than Closely Related Resident Icterids

Environments undergo short-term and long-term changes due to natural or human-induced events. Animals differ in their ability to cope with such changes which can be related to their ecology. Changes in the environment often elicit avoidance reactions (neophobia) which protect animals from dangerous situations but can also inhibit exploration and familiarization with novel situations and thus, learning about new resources. Studies investigating the relationship between a species’ ecology and its neophobia have so far been restricted to comparing only a few species and mainly in captivity. The current study investigated neophobia reactions to experimentally-induced changes in the natural environment of six closely-related blackbird species (Icteridae), including two species represented by two distinct populations. For analyses, neophobic reactions (difference in number of birds feeding and time spent feeding with and without novel objects) were related to several measures of ecological plasticity and the migratory strategy (resident or migratory) of the population. Phylogenetic relationships were incorporated into the analysis. The degree of neophobia was related to migratory strategy with migrants expressing much higher neophobia (fewer birds feeding and for a shorter time with objects present) than residents. Furthermore, neophobia showed a relationship to diet breadth with fewer individuals of diet generalists than specialists returning when objects were present supporting the dangerous niche hypothesis. Residents may have evolved lower neophobia as costs of missing out on opportunities may be higher for residents than migrants as the former are restricted to a smaller area. Lower neophobia allows them approaching changes in the environment (e.g. novel objects) quickly, thereby securing access to resources. Additionally, residents have a greater familiarity with similar situations in the area than migrants and the latter may, therefore, initially stay behind resident species.     

Responses of domestic chicks to novel food as a function of sex,strain and previous experience

Food neophobia or a hesitancy to feed on unfamiliar food was examined in domestic chicks as a function of genetic and experiential factors associated with differential responsiveness to novelty. Chicks were presented with novel blue mash following a regime of food withdrawal and return. ‘S’ line chicks (light hybrid) fed significantly sooner, longer and more than did medium-hybrid ‘T’ line birds and females accepted novel food more readily than males. Food neophobia was significantly reduced by previous exposure to a variety of coloured foods and by enrichment of the home environment with assorted novel objects. The results are discussed in terms of the relationship between food neophobia, general responsiveness to novelty and underlying levels of fearfulness.     

Spatial Neophilia and Spatial Neophobia in Resident and Migratory Warblers (Sylvia)

Residents and migrants use their environment very differently – the former remain in a given habitat throughout the year, whereas the latter are repeatedly confronted with unfamiliar environments. The difference in ecology may influence decision-making processes whether, when and to which extent to explore an unfamiliar environment. We have investigated spatial neophobia and spatial neophilia – two important novelty reactions that may underlie decision-making – in two closely related warbler species, the resident Sardinian warbler and the migratory garden warbler. Individuals of both species could access an unfamiliar room from a familiar cage. We assessed the conflict between the motivation to enter the novel room (spatial neophilia) and the motivation to avoid it (spatial neophobia) as the frequency and duration of perching on the dowel in the cage, which led to the unfamiliar room before entering it. Furthermore, we measured the latency to enter the novel room and compared the number of individuals of each species entering the room. The combination of the parameters measured allowed assessing the degree of both neophobia and neophilia. Finally, the time spent on each branch in the novel room was taken as a measure for spatial exploration. The migrants perched less often and spent less time on the dowel leading to the room, and entered the novel room quicker than the residents. Additionally, more migrants than residents entered the room. The migrants' decision to enter the novel room can best be explained with a combination of low spatial neophobia coupled with high spatial neophilia, whereas the residents' decision-making is best explained with high spatial neophobia coupled with high spatial neophilia. The differences in neophobia support the migrant-neophobia hypothesis. When in the room, the migrants spent less time on each branch than the residents, possibly indicating that the former collect less spatial information than the latter.     

The Significance of Ecological Factors for Exploration and Neophobia in Parrots

Exploratory behaviour plays an important role in most animals for gathering information about their environment. If it constitutes an adaptation to different environmental conditions exploratory behaviour should differ between species. This has been tested with several hypotheses.
Sixty-one parrot species (Psittacidae) from eight tribes with different diets and habitat preferences were investigated in aviaries. Two tests were carried out. First, a novel object (wooden ring) in the familiar aviary was presented on two test days in the exploration test. Latencies until first contact with the object and the duration of exploration were recorded. Secondly, in the neophobia test, novel objects were placed beside the feeding dish and latencies until first food intake were recorded.
The exploration and neophobia data were related to 12 (13) ecological variables using multiple regression analyses. Phylogenetic relationships were considered.
Species that inhabit complex habitats, such as forest edges, or that feed on buds or species from islands showed the shortest latencies in the exploration test. In contrast, long latencies were related to a diet including a great amount of seeds and/or flowers. The longest duration of exploration occurred in species eating nuts or originating from islands, whereas short durations were related to feeding on seeds. Neophobia was positively related to a diet consisting of insects, and negatively to a diet of leaves. There was no relationship between measures of exploration and neophobia. Exploration and neophobia seem to be tightly related to the ecology of a species.     

Explaining the Paradox of Neophobic Explorers: The Social Information Hypothesis

How animals react to novel food and objects is commonly thought of as a crucial step toward innovations. One would therefore expect innovative species to be attracted to novelty and benefit from a combination of low neophobia and a high motivation to explore. Here we draw attention to the innovation paradox: the most innovative species tend to show neophobic reactions when confronted with novel objects or food, but can use social cues to overcome their initial neophobia. Work on novelty response has highlighted the role of ecological factors as determinants of neophobia and exploration tendency. We examine social influences on novelty response and present the idea that social factors enable some species to maintain the paradoxical combination of high neophobia and high exploration tendency. We compare primates with other species, to assess the extent to which primates are unusual. We review empirical studies that show how intrinsic neophobia is generally overcome by social facilitation and social information, i.e., the presence of experts, especially in species with slow life history, probably because social information reduces risk. We also briefly discuss the role of environmental risk in reducing intrinsic neophobia, in particular its absence in captivity. We draw attention to a strong neophobia-reducing effect of being in captivity, due to humans acting as sources of social information. We propose that species showing the paradoxical combination of strong neophobia and strong exploration tendency use social information to select aspects of the environment worth exploring. The social information hypothesis thus offers an explanation for the paradox of neophobic explorers.     

Personality traits and behavioral syndromes in differently urbanized populations of house sparrows (Passer domesticus)

Urbanization creates novel environments for wild animals where selection pressures may differ drastically from those in natural habitats. Adaptation to urban life involves changes in various traits, including behavior. Behavioral traits often vary consistently among individuals, and these so-called personality traits can be correlated with each other, forming behavioral syndromes. Despite their adaptive significance and potential to act as constraints, little is known about the role of animal personality and behavioral syndromes in animals' adaptation to urban habitats. In this study we tested whether differently urbanized habitats select for different personalities and behavioral syndromes by altering the population mean, inter-individual variability, and correlations of personality traits. We captured house sparrows (Passer domesticus) from four different populations along the gradient of urbanization and assessed their behavior in standardized test situations. We found individual consistency in neophobia, risk taking, and activity, constituting three personality axes. On the one hand, urbanization did not consistently affect the mean and variance of these traits, although there were significant differences between some of the populations in food neophobia and risk taking (both in means and variances). On the other hand, both urban and rural birds exhibited a behavioral syndrome including object neophobia, risk taking and activity, whereas food neophobia was part of the syndrome only in rural birds. These results indicate that there are population differences in certain aspects of personality in house sparrows, some of which may be related to habitat urbanization. Our findings suggest that urbanization and/or other population-level habitat differences may not only influence the expression of personality traits but also alter their inter-individual variability and the relationships among them, changing the structure of behavioral syndromes.     

Food Neophobia and Social Learning Opportunities in Great Apes

Finding food resources and maintaining a balanced diet are major concerns for all animals. A compromise between neophobia and neophilia is hypothesised to enable animals to enlarge their diet while limiting the risk of poisoning. However, little is known about how primates respond to novel food items and whether their use is socially transmitted. By comparing how four different species of great apes respond to novel food items, we investigated how differences in physiology (digestive tract size and microbial content), habitats (predictability of food availability), and social systems (group size and composition) affect their response toward novelty. We presented two familiar foods, one novel fruit, four novel aromatic plants from herbal medicine, and kaolin to captive chimpanzees (Pan troglodytes), western gorillas (Gorilla gorilla), Bornean orangutans (Pongo pygmaeus) and Sumatran orangutans (Pongo abelii). We recorded smelling, approach-taste delays, ingestion, interindividual observations, and food transfers with continuous sampling. We found that behaviors differed between the apes: chimpanzees were the most cautious species and observed their conspecifics handling the items more frequently than the other apes. Close observations and food transfers were extremely rare in gorillas in comparison to orangutans and chimpanzees. We suggest that a low neophobia level reflects an adaptive response to digestive physiological features in gorillas and to unpredictable food availability in orangutans. Social interactions appeared to be predominant in chimpanzees and in both orangutan species to overcome food neophobia. They reflect higher social tolerance and more opportunities for social learning and cultural transmission in a feeding context.     

Dominance and neophobia in coyote (Canis latrans) breeding pairs

Previous research suggests that within social animals, subordinate individuals are less neophobic than dominant individuals. We investigated the effect of social status on neophobic responses using 10 captive coyote breeding pairs. Social status was determined from observations of feeding behavior and agonistic interactions during a series of reference trials. Once dominance was established, we surrounded an experimental area with novel stimuli (ropes adorned with interspersed flags) to create a novel context around a familiar food source. Contrary to hypotheses, dominant coyotes were first to feed, showed more interest toward novel stimuli, and eventually crossed the barrier, which subordinates never did. Our results indicate that dominant coyotes are less neophobic of novel settings that contain familiar food than subordinates are. Since a reduction in neophobia can be interpreted as an increase in risk taking, our results support previous observations that dominant (alpha) coyotes take more risks than subordinates. Our results also suggest reasons for differential observations of coyote behavior in the field: artificial selection against bold behavior in populations undergoing predator control.     

Deactivation of dietary wariness through experience of novel food

When a forager encounters an unfamiliar type of food, it mustdecide whether to eat it and risk being poisoned or avoid eatingit and risk forfeiting a potentially valuable resource. Birdstypically respond to such situations with "dietary wariness";they show a transient aversion to approaching new food (neophobia),and many individuals also show a much longer lasting reluctanceto consume the new food (dietary conservatism), even once neophobiahas waned. Very little is known about how these processes, togethertermed "wariness," are controlled. We therefore present a seriesof experiments investigating how wariness of novel foods indomestic chicks, Gallus gallus domesticus, can be deactivatedand reactivated by different experiences of colored foods, varyingin their degree of novelty and palatability. We found that priorexperience of a single novel color of palatable chick crumbswas sufficient to deactivate both neophobia and dietary conservatismof any other novel color of crumbs tested. Relatively littleprior experience of a novel training food was needed to deactivateneophobia, after which the birds would peck at any other novelfood. In contrast, much more extensive experience of eatinga novel training food was needed before the birds would incorporateother novel foods into their diet. Chicks needed direct physicalcontact with the training food before they overcame their warinessto eat another novel food. However, observational learning wassufficient to encourage them to peck at the food (overcomingtheir neophobia). Reinstating wariness was much more easilyachieved than its deactivation. We discuss these surprisingresults in relation to the foraging behavior of wild and domesticbirds.     

Head Colour and Age Relate to Personality Traits in Gouldian Finches

Colour polymorphism is a widespread phenomenon in vertebrates and has often been linked with differences in behaviour such as aggression or boldness, behaviours that are often part of personality traits in monomorphic species. However, up to now, very few studies have looked whether colour morphs just differ in average behaviour or whether specific combinations of behaviours are favoured in relation to a particular morph therefore signalling personality traits. This was tested in the highly social and polymorphic Gouldian finch (Erythrura gouldiae). Tests were performed in pairs of a red‐headed and a black‐headed individual of same sex and size but different age. Capture rank in the aviary as well as latency to feed beside a novel object (neophobia) and to approach and touch a novel object on a perch (neophilia) were measured. For neophobia and neophilia, ranks of latencies calculated across all individuals were used and general and within‐pair comparisons were made. Neophobia and approach neophilia were consistent over time (2 mo), and approach neophilia was positively correlated with capture rank and neophobia indicating that the behaviours are part of personality traits. Moreover, black‐headed birds generally approached the novel object earlier than red‐headed birds. Similarly, within pairs, black‐headed birds were first to approach the novel object and were also captured first. Relative age (younger or older than the partner) was related to neophobia and tactile neophilia ranks; older birds reacted faster. The results indicate that colour morphs not only differ in average behaviour but that specific combinations of behaviours are related to colour morphs suggesting that head colours signal personality traits. Furthermore, depending on age individuals may have different roles in a social network.     

Sex‐Specific Associations Between Nest Defence,Exploration and Breathing Rate in Breeding Pied Flycatchers

Correlations between behavioural traits constrain animals to a limited range of behavioural choices and set a limit to the available variation in behavioural phenotype of a population. These constraints are especially important during stressful situations, potentially limiting the ability to cope with stress appropriately. As yet, very little consideration has been given to a possible role for sexual selection in maintaining differences in behavioural stability within individuals: sex‐specific differences in behavioural correlations have been seldom studied, especially in the wild. In this field study, we investigated associations between neophobia (latency to enter nest in the presence of a novel object), nest defence (response to a model predator) and breathing rate in response to handling in breeding pied flycatchers (Ficedula hypoleuca). We did not find any significant difference in antipredator responses between males and females, which indicates low sexual conflict over parental care in the pied flycatcher. However, females were more neophobic than males, while males and females did not differ in their breathing rates. Further, our study demonstrated a strong positive correlation between nest defence behaviour and neophobia in male, but not in female pied flycatchers. Males that defended their nest more had lower breathing rate and higher latency to resume nestling feeding after encountering a predator decoy than males that mobbed less intensely. We found only weak evidence that nest partners might affect each other's behaviour in these contexts.     

Early experience affects learning performance and neophobia in a cooperatively breeding cichlid

The ability to respond flexibly to environmental challenges, for instance by learning or by responding appropriately to novel stimuli, may be crucial for survival and reproductive success. Experiences made during early ontogeny can shape the degree of behavioural flexibility maintained by individuals during later life. In natural habitats, animals are exposed to a multitude of social and non‐social ecological factors during early ontogeny, but their relative influences on future learning ability and behavioural flexibility are only poorly understood. In the cooperatively breeding cichlid Neolamprologus pulcher, we investigated whether early social and predator experiences shape the learning performance, flexibility, and response to novelty of adults. Fish were reared either with or without parents and helpers and with or without perceived predation risk in a full‐factorial experiment. We investigated the influence of these treatments on learning performance and flexibility in a spatial acquisition and reversal learning task. To test for response to novelty, we performed a neophobia test. We found that fish reared with predator experience, but without the presence of older group members outperformed fish with other rearing backgrounds in reversal learning and that individuals, which had been reared in a socially more complex environment together with older group members responded less neophobic toward a novel object than individuals reared among siblings only. Comparative evidence from fish and rats suggests that these developmental effects may be driven by the cues of safety perceived in the presence of guarding parents.     

Individual and Sex Differences in Habituation and Neophobia in House Sparrows (Passer domesticus)

Heterogeneity in the environment favours foragers that are flexible (phenotypically plastic). However, consistent individual differences in behaviour (personality), such as in risk‐taking, might affect an individual's ability to find food, avoid predators and adjust to new conditions. It is now well known that personalities exist in many taxa, but much less is known about individual variation in plasticity. We measured the tendency to begin foraging across different levels of risk in house sparrows (Passer domesticus), using a behavioural reaction norm framework to simultaneously assess personality and plasticity. We asked whether individuals were consistently different across risk levels, and whether they differed in habituation and neophobia, both of which were treated as cases of plasticity. We found that males habituated more than females by beginning to feed sooner after repeated instances of a human disturbance in the presence of an initially unfamiliar object. Individuals of both sexes also exhibited consistent differences across trials, but did not differ in the rate of habituation beyond the difference between the sexes. When a novel object was placed in the foraging area, both sexes exhibited similar degrees of neophobia by delaying feeding. The magnitude of this change varied among birds, indicating individual differences in neophobia. Our results indicate that both personality and individual variation in plasticity exist but should be treated as independent phenomena. The presence of variation in plasticity implies that the raw material necessary for selection on neophobia exists, and that if heritable, plasticity in risk‐taking across contexts could evolve.     

Do neophobia and dietary wariness explain ecological flexibility? An analysis with two seed‐eating birds of contrasting habits

The neophobia threshold hypothesis (NTH) suggests that the acquisition and maintenance of a high behavioral and ecological flexibility in the evolutionary and adaptive history of a species is the consequence of lower levels of neophobia towards new micro‐habitats and of dietary wariness of novel foods. To test this idea we assessed the degree of neophobia and dietary wariness in two seed‐eating bird species with contrasting degrees of ecological flexibility that inhabit the central Monte desert (Argentina): a grass‐seed specialist, the many‐colored chaco‐finch, and a generalist feeder, the rufous‐collared sparrow. We expected that both species would exhibit neophobia and wariness when faced with new foraging opportunities but that the rufous‐collared sparrow would be less neophobic and less wary than the specialized many‐colored chaco‐finch. Experimental indicators of neophobia and dietary wariness included willingness to eat near novel objects and willingness to eat novel seeds, respectively. Both species showed similar levels of reluctance to novelty, although the sparrow could be slightly more reluctant than the finch. Contrary to our predictions, the sparrow was neither less hesitant nor faster or greedier than the finch. This experimental evidence does not support a negative relationship between neophobia/wariness and ecological flexibility in these two seed‐eating birds and it coincides with the growing evidence that challenges the NTH. Some of our results provide support for the dangerous niche hypothesis, especially as the rufous‐collared sparrow, that feeds on more diverse and potentially dangerous food, showed higher levels of neophobia in some cases. Although the idea of neophobia and wariness being plausible causes of ecological specialization sounds attractive, the current situation calls for further research so that the causes of ecological flexibility in granivorous birds can be better understood.     

Maternal presence and environmental enrichment affect food neophobia of piglets

Young omnivores show food neophobia in order to avoid the potential harmful effects of ingesting unfamiliar food items. We investigated whether the presence of the mother and an enriched rearing environment would reduce food neophobia in piglets. A mother may provide information on suitable food types to include in the diet, whereas an enriched environment may stimulate behavioural development and reduce reactivity towards novel stimuli (including food). Five barren-reared or enriched-reared piglets per litter were exposed to two novel food items in the presence, and the other five per litter in the absence, of the mother in a 7 min test. Maternal presence reduced food neophobia profoundly as reflected in a reduced latency to touching the food, a higher proportion of piglets sampling the two different food items and a higher intake. Latency to touch the food, however, was affected by maternal presence more strongly for barren-reared piglets than for enriched-reared piglets, and in the absence of the sow, consumption of one novel food type and time spent in the feeding area were higher for enriched-reared piglets.Environmental enrichment does have the potential to reduce food neophobia, but the presence of the mother during the encounter with novel food seems more efficient in decreasing food neophobia of piglets.     

Behavioural and hormonal effects of social isolation and neophobia in a gregarious bird species, the European starling (Sturnus vulgaris)

Separating gregarious individuals from their group members often results in behavioural and physiological changes, like increased levels of corticosterone. Testosterone and corticosterone, in particular, have been implicated in the response of mammals to novelty. Data in birds are, however, rare. The presence or absence of group members may also influence an individual's response to novel stimuli. We assessed the behaviour and hormonal response of European starlings (Sturnus vulgaris) to a novel object in two different situations and seasons: each starling was tested when separated and when in contact with its group members in May/June (breeding season) and again in September/October (non-breeding season). Starlings are gregarious throughout the year, but as foraging flocks are small during the breeding season and large during the non-breeding season, we assumed that non-breeding starlings would be more affected by social isolation. Overall, starlings had higher levels of corticosterone, lost more body mass, and were more active when they were separated from their group. Isolated individuals, however, did not show a greater neophobic response than individuals in the presence of their group members in either season. Circulating levels of testosterone and corticosterone were higher after a test with novel object than after a test with only the familiar feeding dish in both sexes and seasons. However, control tests for handling effects confirmed only the increase in testosterone. Our study shows that social isolation is stressful for unrelated and unpaired members of a wild flocking bird species and demonstrates that novelty can lead to a rise in testosterone in birds.     

Predator-induced neophobia in juvenile cichlids

Predation is an important but often fluctuating selection factor for prey animals. Accordingly, individuals plastically adopt antipredator strategies in response to current predation risk. Recently, it was proposed that predation risk also plastically induces neophobia (an antipredator response towards novel cues). Previous studies, however, do not allow a differentiation between general neophobia and sensory channel-specific neophobic responses. Therefore, we tested the neophobia hypothesis focusing on adjustment in shoaling behavior in response to a novel cue addressing a different sensory channel than the one from which predation risk was initially perceived. From hatching onwards, juveniles of the cichlid Pelvicachromis taeniatus were exposed to different chemical cues in a split-clutch design: conspecific alarm cues which signal predation risk and heterospecific alarm cues or distilled water as controls. At 2 months of age, their shoaling behavior was examined prior and subsequent to a tactical disturbance cue. We found that fish previously exposed to predation risk formed more compact shoals relative to the control groups in response to the novel disturbance cue. Moreover, the relationship between shoal density and shoal homogeneity was also affected by experienced predation risk. Our findings indicate predator-induced, increased cross-sensory sensitivity towards novel cues making neophobia an effective antipredator mechanism.     

Cautious Crows: Neophobia in Torresian Crows (Corvus orru) Compared with Three Other Corvoids in Suburban Australia

Corvids (Family: CORVIDAE) are a clade of some 120 species widespread throughout much of the world that have attracted the interest of researchers due to their impressive cognitive abilities. The group is, however, also generally described as neophobic, a trait that increases the difficulty of undertaking such research. In Australia, Torresian crows (Corvus orru) have, like corvid species worldwide, thrived in urban environments, sharing this habitat with a number of other corvoid (Superfamily: CORVOIDEA) species. While each of these species has successfully colonised urban areas, the extent to which neophobia is present is not known. This study empirically tested the extent to which neophobia is exhibited in wild urban Torresian crows by measuring the delaying effect of a novel object to obtaining food and any changes in neophobic behaviours displayed. This was then compared with the other urban corvoid species that inhabit similar niches. This study confirmed that Torresian crows are significantly wary of a novel objects, displaying more neophobic behaviours and taking longer to attain the food. Crow behaviour provided evidence in support of both the dangerous niche hypothesis and the two‐factor model of neophobia and neophilia. Crows also displayed these behaviours to a significantly greater extent than the three other corvoids studied. However, the individual variation in crow behaviours when exposed to a novel object was extensive. This variation may be attributed to differing behavioural types between individuals, or different experiences with novel objects or humans in the bird's past.     

COMPARATIVE METHODS AT THE SPECIES LEVEL: GEOGRAPHIC VARIATION IN MORPHOLOGY AND GROUP SIZE IN GREY-CROWNED BABBLERS (POMATOSTOMUS TEMPORALIS)

We show that a new comparative method that sheds light on evolutionary trends among species may also illuminate trends within species. This finding comes from a phylogenetic autocorrelation analysis of morphological traits among individuals sampled from ten populations of a cooperatively breeding songbird, the Grey-crowned Babbler (Pomatostomus temporalis). Highly variable mitochondrial DNA (mtDNA) sequences from both the eastern (Pomatostomus temporalis temporalis) and western (Pomatostomus temporalis rubeculus) lineages were used to define genetic distances among 120 individuals and to estimate correlations among individuals in wing length, tarsus length, and body weight via an intraspecific weighting matrix. The autoregressive model effectively removed intraspecific correlations for all three morphological variables, and the proportion of the total phenotypic variance due to genealogical relationships varied from 0.68 (weight) to 0.23 (tarsus). The analysis revealed correlations among the specific components of traits, in which none were previously detected (type-I error) and diminished correlations that appeared strong when phylogeny was ignored. Group size was the only trait for which the autoregressive model failed to remove intraspecific correlations, a result likely due to the plasticity, convergence, and clinal variation in this trait in both the eastern and western lineages. The autocorrelation analysis weakened significant negative correlations between group size and total values for wing length and body weight, but the interpretation of this result depends on the adaptive significance ascribed to the “phylogenetic component” of trait values removed by the analysis. Comparative methods employing distance matrices are one useful way of summarizing the pattern of nonhierarchical relationships among conspecific individuals (“tokogenetic” relationships, sensu Hennig).