@misc{allee1951cooperation1,
    author = "Allee, W. C",
    title = "Cooperation Among Animals with Human Implications",
    year = "1951",
    howpublished = "New York, Schuman, 233 p.; [Revised Edition of Social Life of Animals, Norton, New York, 1938]",
    note = "talkorigins\_source = {true}; raw\_reference = {Allee, W. C., 1951, Cooperation Among Animals with Human Implications: New York, Schuman, 233 p.; [Revised Edition of Social Life of Animals, Norton, New York, 1938].}"
}

@book{doi10129879780300188479022,
    author = "Odum, Eugene P.",
    title = "Fundamentals of Ecology (1953)",
    year = "1955",
    booktitle = "Yale University Press eBooks",
    abstract = "Preface. Eugene P. Odum and Gary W. Barrett. 1. The Scope of Ecology. 2. The Ecosystem. 3. Energy in Ecological Systems. 4. Biogeochemical Cycles. 5. Limiting and Regulatory Factors. 6. Population Ecology. 7. Community Ecology. 8. Ecosystem Development. 9. Landscape Ecology. 10. Regional Ecology: Major Ecosystem Types and Biomes. 11. Global Ecology. 12. Statistical Thinking for Students of Ecology. Glossary. References. Index.",
    url = "https://doi.org/10.12987/9780300188479-022",
    doi = "10.12987/9780300188479-022",
    openalex = "W2015345446"
}

@incollection{bull1957behavior4,
    author = "Bull, H. O",
    editor = "Brown, M. E.",
    title = "Behavior: Conditioned Response",
    year = "1957",
    booktitle = "The Physiology of Fishes",
    publisher = "New York, Academic Press, v. II, p. 211-228",
    note = "talkorigins\_source = {true}; raw\_reference = {Bull, H. O., 1957, Behavior: Conditioned Response, in Brown, M. E., ed., The Physiology of Fishes: New York, Academic Press, v. II, p. 211-228.}"
}

@incollection{carpenter1958territoriality5,
    author = "Carpenter, C. R",
    editor = "Roe, A. and Simpson, G. G.",
    title = "Territoriality: A Review of Concepts and Problems",
    year = "1958",
    booktitle = "Behavior and Evolution",
    publisher = "New Haven, Yale University Press, p. 224-250; 537 pp",
    note = "talkorigins\_source = {true}; raw\_reference = {Carpenter, C. R., 1958, Territoriality: A Review of Concepts and Problems, in Roe, A., and Simpson, G. G., eds., Behavior and Evolution: New Haven, Yale University Press, p. 224-250; 537 pp.}"
}

@misc{ardrey1966the2,
    author = "Ardrey, R",
    title = "The Territorial Imperative",
    year = "1966",
    howpublished = "New York, Atheneum, 390 p",
    note = "talkorigins\_source = {true}; raw\_reference = {Ardrey, R., 1966, The Territorial Imperative: New York, Atheneum, 390 p.}"
}

@article{doi104319lo19721760805,
    author = "Frost, Bruch W.",
    title = "EFFECTS OF SIZE AND CONCENTRATION OF FOOD PARTICLES ON THE FEEDING BEHAVIOR OF THE MARINE PLANKTONIC COPEPOD CALANUS PACIFICUS 1",
    year = "1972",
    journal = "Limnology and Oceanography",
    abstract = "When adult females of Calanus pacificus are fed on monospecific cultures of centric diatoms which grow as single cells, a predictive relationship is found between feeding behavior of the copepods and size and concentration of food particles. Ingestion rate of copepods increases linearly with cell concentration up to a maximal rate. This maximal ingestion rate, expressed as carbon, is the same for copepods feeding on diatoms ranging in diameter from 11–87 µ. As the size of food particles increases, the carbon concentration at which this ingestion rate is achieved decreases. Thus females of C. pacificus can obtain their maximal daily ration at relatively low carbon concentrations of large cells.",
    url = "https://doi.org/10.4319/lo.1972.17.6.0805",
    doi = "10.4319/lo.1972.17.6.0805",
    openalex = "W2095101020",
    references = "doi104039entm9745fv"
}

@book{openalexw1532540194,
    author = "Krebs, Charles J.",
    title = "Ecology: The Experimental Analysis of Distribution and Abundance",
    year = "1973",
    abstract = {Charles Krebs' best-selling majors-level text approaches ecology as a series of problems that are best understood by evaluating empirical evidence through data analysis and application of quantitative reasoning. No other text presents analytical, quantitative, and statistical ecological information in an equally accessible style for students. Reflecting the way ecologists actually practice, the new edition emphasizes the role of experiments in testing ecological ideas and discusses many contemporary and controversial problems related to distribution and abundance. Ecology: The Experimental Analysis of Distribution and Abundance, Sixth Edition builds on a clear writing style, historical perspective, and emphasis on data analysis with an updated, reorganized discussion of key topics and two new chapters on climate change and animal behavior. Key concepts and key terms are now included at the beginning of each chapter to help students focus on what is most important within each chapter, mathematical analyses are broken down step by step in a new feature called "Working with the Data," concepts are reinforced throughout the text with examples from the literature, and end-of-chapter questions and problems emphasize application},
    openalex = "W1532540194"
}

@misc{farlow1976a7,
    author = "Farlow, J. O",
    title = "A consideration of the trophic dynamics of a Late Cretaceous large- dinosaur community (Oldman Formation)",
    year = "1976",
    howpublished = "Ecology, v. 57, p. 841-857",
    note = "talkorigins\_source = {true}; raw\_reference = {Farlow, J. O., 1976, A consideration of the trophic dynamics of a Late Cretaceous large- dinosaur community (Oldman Formation): Ecology, v. 57, p. 841-857.}"
}

@misc{farlow1976speculations6,
    author = "Farlow, J. O",
    title = "Speculations about the diet and foraging behavior of large carnivorous dinosaurs",
    year = "1976",
    howpublished = "American Midland Naturalist, v. 95, p. 186-191",
    note = "talkorigins\_source = {true}; raw\_reference = {Farlow, J. O., 1976, Speculations about the diet and foraging behavior of large carnivorous dinosaurs: American Midland Naturalist, v. 95, p. 186-191.}"
}

@misc{gruber1977approaches8,
    author = "Gruber, S. H. and Myrberg, A. A. J",
    title = "Approaches to the study of the behavior of sharks",
    year = "1977",
    howpublished = "American Zoologist, v. 17, p. 471-486",
    note = "talkorigins\_source = {true}; raw\_reference = {Gruber, S. H., and Myrberg, A. A. J., 1977, Approaches to the study of the behavior of sharks: American Zoologist, v. 17, p. 471-486.}"
}

@article{hopson1977relative9,
    author = "Hopson, J. A",
    title = "Relative brain size and behavior in archosaurian reptiles",
    year = "1977",
    journal = "Annual Review of Ecology and Systematics, v. 8, p. 429-448",
    note = "talkorigins\_source = {true}; raw\_reference = {Hopson, J. A., 1977, Relative brain size and behavior in archosaurian reptiles: Annual Review of Ecology and Systematics, v. 8, p. 429-448.}"
}

@article{doi101126science21044731041,
    author = "Sih, Andrew",
    title = "Optimal Behavior: Can Foragers Balance Two Conflicting Demands?",
    year = "1980",
    journal = "Science",
    abstract = "According to much current theory, organisms should be able to balance the conflicting demands of the need to feed efficiently and the need to avoid preadtors while feeding. In an experimental conflict situation, it was possible to evaluate the relative fitnesses associated with the available choices and to compare the observed behaviors with predictions derived from fitness considerations. A backswimmer, Notonecta hoffmanni, was capable of balancing these two conflicting factors adaptively.",
    url = "https://doi.org/10.1126/science.210.4473.1041",
    doi = "10.1126/science.210.4473.1041",
    openalex = "W2024142792",
    references = "doi101016004058097690040x, doi1010160040580977900466, doi101038275642a0, doi101086282454, doi101086282885, doi101086409852, doi101146annureves07110176001315, doi1023071934203, doi104319lo19762160804, openalexw1487200503"
}

@book{auffenburg1981the3,
    author = "Auffenburg, W",
    title = "The Behavorial Ecology of the Komodo Monitor",
    year = "1981",
    publisher = "Gainesville, Florida, University of Florida Presses",
    note = "talkorigins\_source = {true}; raw\_reference = {Auffenburg, W., 1981, The Behavorial Ecology of the Komodo Monitor: Gainesville, Florida, University of Florida Presses.}"
}

@article{davies1981behavioral,
    author = "DAVIES, N. B.",
    title = "Behavioral Ecology",
    year = "1981",
    journal = "Science",
    url = "https://doi.org/10.1126/science.211.4488.1341",
    doi = "10.1126/science.211.4488.1341",
    number = "4488",
    pages = "1341-1341",
    volume = "211"
}

@article{doi101007bf00005175,
    author = "McKaye, Kenneth Rober",
    title = "Ecology and breeding behavior of a cichlid fish, Cyrtocara eucinostomus, on a large lek in Lake Malawi, Africa",
    year = "1983",
    journal = "Environmental Biology of Fishes",
    url = "https://doi.org/10.1007/bf00005175",
    doi = "10.1007/bf00005175",
    openalex = "W2087031354",
    references = "doi101073pnas77116937, doi101126science1188360, doi1023071443541, doi1023072063069, doi1023072576242, doi1023072874, doi1023073421, doi1023074335, doi1043249781315129266, openalexw2060759128"
}

@article{doi1023074788,
    author = "Harrington, R. and Slansky, Frank and Rodríguez, J. G.",
    title = "Nutritional Ecology of Insects, Mites, Spiders and Related Invertebrates",
    year = "1988",
    journal = "Journal of Animal Ecology",
    abstract = "Nutritional Ecology of Forb Foliage-Chewing Insects Nutritional Ecology of Insect Folivores of Woody Plants: Nitrogen, Water, Fiber and Mineral Considerations Nutritional Ecology of Grass Foliage-Chewing Insects Nutritional Ecology of Phytophagous Mites Nutritional Ecology of Lichen/Moss Arthropods Nutritional Ecology of Arthropod Gall-Makers Nutritional Ecology of Bruchid Beetles Nutritional Ecology of Seed-Sucking Insects Nutritional Ecology of Stored-Product Insects Nutritional Ecology of Stored-Product and House Dust Mites Ecology of Nectar and Pollen Feeding in Lepidoptera.",
    url = "https://doi.org/10.2307/4788",
    doi = "10.2307/4788",
    openalex = "W2068793744"
}

@article{doi101139z90092,
    author = "Lima, Steven L. and Dill, Lawrence M.",
    title = "Behavioral decisions made under the risk of predation: a review and prospectus",
    year = "1990",
    journal = "Canadian Journal of Zoology",
    abstract = "Predation has long been implicated as a major selective force in the evolution of several morphological and behavioral characteristics of animals. The importance of predation during evolutionary time is clear, but growing evidence suggests that animals also have the ability to assess and behaviorally influence their risk of being preyed upon in ecological time (i.e., during their lifetime). We develop an abstraction of the predation process in which several components of predation risk are identified. A review of the literature indicates that an animal's ability to assess and behaviorally control one or more of these components strongly influences decision making in feeding animals, as well as in animals deciding when and how to escape predators, when and how to be social, or even, for fishes, when and how to breathe air. This review also reveals that such decision making reflects apparent trade-offs between the risk of predation and the benefits to be gained from engaging in a given activity. Despite this body of evidence, several areas in the study of animal behavior have received little or no attention from a predation perspective. We identify several such areas, the most important of which is that dealing with animal reproduction. Much work also remains regarding the precise nature of the risk of predation and how it is actually perceived by animals, and the extent to which they can behaviorally control their risk of predation. Mathematical models will likely play a major role in future work, and we suggest that modelers strive to consider the potential complexity in behavioral responses to predation risk. Overall, since virtually every animal is potential prey for others, research that seriously considers the influence of predation risk will provide significant insight into the nature of animal behavior.",
    url = "https://doi.org/10.1139/z90-092",
    doi = "10.1139/z90-092",
    openalex = "W2165111556",
    references = "doi101007bf00346972, doi101007bf00395696, doi101016016953479190210o, doi101016s0065345408601928, doi101086409052, doi101126science21044731041, doi101126science7466396, doi101146annurevecolsys151393, doi101146annurevento43163, doi101146annureves02110171002101, doi1023071937508, doi104039ent912935"
}

@incollection{crossref1991behavioral,
    title = "Behavioral Ecology",
    year = "1991",
    booktitle = "Techniques in the Behavioral and Neural Sciences",
    url = "https://doi.org/10.1016/b978-0-444-81251-3.50011-4",
    doi = "10.1016/b978-0-444-81251-3.50011-4",
    pages = "117-153"
}

@article{doi101146annureven37010192001041,
    author = "Vet, L.E.M. and Dicke, Marcel",
    title = "Ecology of Infochemical Use by Natural Enemies in a Tritrophic Context",
    year = "1992",
    journal = "Annual Review of Entomology",
    abstract = "Parasitoids and predators of herbivores have evolved and function within a multitrophic context.Consequently, their physiology and behavior are influenced by elements from other trophic levels such as their herbivore victim (second trophic level) and its plant food (first trophic level) (126).Natural enemies base their foraging decisions on information from these different trophic levels, and chemical information plays an important role.This review is restricted to the ecology of chemical information from the first and second trophic levels.The importance of so-called infochemicals, a subcategory of semiochemicals, in foraging by parasitoids and predators has been well documented (e.g.reviewed in 31,78, 111,183,185), and we do not intend repeat the details.But because of a lack of testable hypotheses, all this research is conducted rather haphazardly: the total puzzle of infochemical use has not been solved for any natural enemy species.Here we approach the use of infochemicals by natural enemies from an evolutionary and ecological standpoint.Our basic concept is that information from the first and second trophic levels differs in availability and in reliability, a difference that shapes the way infochemicals are used by a species.We generate hypotheses on (a)",
    url = "https://doi.org/10.1146/annurev.en.37.010192.001041",
    doi = "10.1146/annurev.en.37.010192.001041",
    openalex = "W2163028358",
    references = "doi101086282454, doi101126science185414527, doi101126science2114485887"
}

@article{doi1023071468026,
    author = "Poff, N. LeRoy",
    title = "Landscape Filters and Species Traits: Towards Mechanistic Understanding and Prediction in Stream Ecology",
    year = "1997",
    journal = "Journal of the North American Benthological Society",
    abstract = {A heuristic framework for understanding and predicting the distribution and categorical abundance of species in stream communities is presented. The framework requires that species be described in terms of their functional relationships to habitat selective forces or their surrogates, which constitute "filters" occurring at hierarchical landscape scales (ranging from microhabitats to watersheds or basins). Large-scale filters are viewed as causative or mechanistic agents that constrain expression of local selective forces or biotic potential at lower scales. To join a local community, species in a regional pool must possess appropriate functional attributes (species traits) to "pass" through the nested filters. Biotic interactions are also a potential filter on local community composition, and they are invoked at the lower hierarchical levels, after species have passed through the physicochemical habitat filters. Potential landscape filters and their associated selective properties are identified, as are prospective species traits (for invertebrates and fish) that correspond with filters. A categorical niche model is used to illustrate how relative abundances of species in local communities might be predicted from habitat data collected at different scales. The framework emphasizes a biologically based approach to understanding and predicting species distribution and abundance and local community composition by explicitly considering environmental constraints imposed at different scales. As such, it can complement non-mechanistic, correlative approaches to community prediction that often lack generality. Operationalizing the framework will require additional research to specify more clearly 1) the degree to which habitat features at different scales are linked functionally or statistically, 2) what species traits are possessed by strongly interactive species (e.g., keystones) and which habitat filters most strongly constrain the distribution of these species, and 3) the functional significance of a range of species traits and the extent to which these traits are correlated and hence respond in concert to the presence, or modification, of a particular filter. Multi-scale, mechanistic understanding of species-environment relations will likely contribute to better predictions about large scale problems, such as the establishment and spread of exotic species or alterations in community composition with changing land use or climate.},
    url = "https://doi.org/10.2307/1468026",
    doi = "10.2307/1468026",
    openalex = "W2030196717",
    references = "doi101007978146124018114, doi101086285270, doi101139f80017, doi1023071942327, doi1023075403"
}

@incollection{crossref1998behavioral,
    title = "Behavioral Ecology",
    year = "1998",
    booktitle = "Comparative Psychology",
    url = "https://doi.org/10.4324/9780203826492-17",
    doi = "10.4324/9780203826492-17",
    pages = "97-103"
}

@article{doi10108010420949809386437,
    author = "Miller, William L. and Vokes, Emily H.",
    title = "Large Phymatoderma in pliocene slope deposits, northwestern Ecuador: Associated ichnofauna, fabrication, and behavioral ecology",
    year = "1998",
    journal = "Ichnos/Ichnos : an international journal for plant and animal traces",
    abstract = "Phytnatoderma is a complex, branching biogenic structure that is seldom mentioned in the ichnologic literature, although it may have been misidentified recently as “Zonarites” and large pellet‐filled “Chondrites”. The ichnogenus is usually interpreted as the product of subsurface deposit‐feeding animals. Well‐preserved examples from Pliocene slope mudstone in Ecuador are assignable to P. granulatum (Schlotheim, 1822). These structures are typically 30–40 cm in overall diameter; consist of outward‐branching, horizontal to subhorizontal, overlapping tunnels grouped in bundles; and have tunnel fills consisting of pelleted volcanic ash. The Ecuadorian Phymatoderma occur with a moderately rich ichnoassemblage that includes Alcyonidiopsis ichnosp., Chondrites intricatus, several kinds of Phycodes‐like structures, Phycosiphon incertum, Planolites ichnosp., Rorschachichnus amoeba, Scolicia ichnosp., Taenidium barretti, and Zoophycos ichnosp. Benthic foraminiferids and geologic context point to deposition on the upper trench slope, ca. 3–4 Ma. Ecuadorian Phymatoderma are large, structurally intricate burrow systems that not only represent surface deposit feeding and below‐surface pellet stowage/food caching, but also behavior that apparently included revisiting runnels to utilize the pellets for food, in response to a variable trophic regime. Together with other morphologically intricate, ethologically complex marine ichnogenera (e.g., Zoophycos, Chondrites, Paleodictyon), Phymatoderma deserves special attention as a complex animal artefact, rather than being considered merely a snapshot of a single or dominant behavior. Such complicated structures appear to record deliberate activity to restructure habitats, modulate natural disturbances, and to control trophic resources.",
    url = "https://doi.org/10.1080/10420949809386437",
    doi = "10.1080/10420949809386437",
    openalex = "W2076849361",
    references = "doi101007978146124018114, doi1010079783642748646, doi101017cbo9781139163637, doi101038141548c0, doi101111j136531211991tb00851x, doi1023073545850, doi105860choice293872, doi105860choice295709, openalexw1486903121, openalexw2912219260"
}

@article{doi1023071383287,
    author = "Brown, J. S. and Laundre, J. W. and Gurung, M. B.",
    title = "The Ecology of Fear: Optimal Foraging, Game Theory, and Trophic Interactions",
    year = "1999",
    journal = "Journal of Mammalogy",
    abstract = "Mammalian predator-prey systems are behaviorally sophisticated games of stealth and fear. But, traditional mass-action models of predator prey dynamics treat individuals as behaviorally unresponsive “molecules” in Brownian motion. Foraging theory should provide the conceptual framework to envision the interaction. But, current models of predator feeding behavior generally envision a clever predator consuming large numbers of sessile and behaviorally inert prey (e.g., kangaroo rats, Dipodomys, collecting seeds from food patches). Here, we extend foraging theory to consider a predator-prey game of stealth and fear and then embed this game into the modeling of predator-prey population dynamics. The melding of the prey and predator's optimal behaviors with their population and community-level consequences constitutes the ecology of fear. The ecology of fear identifies the endpoints of a continuum of N-driven (population size) versus μ-driven (fear) systems. In N-driven systems, the major direct dynamical feedback involves predators killing prey, whereas μ-driven systems involve the indirect effects from changes in fear levels and prey catchability. In μ-driven systems, prey respond to predators by becoming more vigilant or by moving away from suspected predators. In this way, a predator (e.g., mountain lion, Puma concolor) depletes a food patch (e.g., local herd of mule deer, Odocoileus hemionus) by frightening prey rather than by actually killing prey. Behavior buffers the system: a reduction in predator numbers should rapidly engender less vigilant and more catchable prey. The ecology of fear explains why big fierce carnivores should be and can be rare. In carnivore systems, ignore the behavioral game at one's peril.",
    url = "https://doi.org/10.2307/1383287",
    doi = "10.2307/1383287",
    openalex = "W2005745339",
    references = "doi101007bf00395696, doi1010160040580977900429, doi101016s0065250408602883, doi101086282272, doi101086282454, doi101086282455, doi101126science21044731041, doi1023073280305, doi104039entm9745fv, openalexw1484524608"
}

@article{doi101111j001438202003tb00285x,
    author = "Blomberg, Simon P. and Garland, Theodore and Ives, Anthony R.",
    title = "TESTING FOR PHYLOGENETIC SIGNAL IN COMPARATIVE DATA: BEHAVIORAL TRAITS ARE MORE LABILE",
    year = "2003",
    journal = "Evolution",
    abstract = "The primary rationale for the use of phylogenetically based statistical methods is that phylogenetic signal, the tendency for related species to resemble each other, is ubiquitous. Whether this assertion is true for a given trait in a given lineage is an empirical question, but general tools for detecting and quantifying phylogenetic signal are inadequately developed. We present new methods for continuous-valued characters that can be implemented with either phylogenetically independent contrasts or generalized least-squares models. First, a simple randomization procedure allows one to test the null hypothesis of no pattern of similarity among relatives. The test demonstrates correct Type I error rate at a nominal alpha = 0.05 and good power (0.8) for simulated datasets with 20 or more species. Second, we derive a descriptive statistic, K, which allows valid comparisons of the amount of phylogenetic signal across traits and trees. Third, we provide two biologically motivated branch-length transformations, one based on the Ornstein-Uhlenbeck (OU) model of stabilizing selection, the other based on a new model in which character evolution can accelerate or decelerate (ACDC) in rate (e.g., as may occur during or after an adaptive radiation). Maximum likelihood estimation of the OU (d) and ACDC (g) parameters can serve as tests for phylogenetic signal because an estimate of d or g near zero implies that a phylogeny with little hierarchical structure (a star) offers a good fit to the data. Transformations that improve the fit of a tree to comparative data will increase power to detect phylogenetic signal and may also be preferable for further comparative analyses, such as of correlated character evolution. Application of the methods to data from the literature revealed that, for trees with 20 or more species, 92\% of traits exhibited significant phylogenetic signal (randomization test), including behavioral and ecological ones that are thought to be relatively evolutionarily malleable (e.g., highly adaptive) and/or subject to relatively strong environmental (nongenetic) effects or high levels of measurement error. Irrespective of sample size, most traits (but not body size, on average) showed less signal than expected given the topology, branch lengths, and a Brownian motion model of evolution (i.e., K was less than one), which may be attributed to adaptation and/or measurement error in the broad sense (including errors in estimates of phenotypes, branch lengths, and topology). Analysis of variance of log K for all 121 traits (from 35 trees) indicated that behavioral traits exhibit lower signal than body size, morphological, life-history, or physiological traits. In addition, physiological traits (corrected for body size) showed less signal than did body size itself. For trees with 20 or more species, the estimated OU (25\% of traits) and/or ACDC (40\%) transformation parameter differed significantly from both zero and unity, indicating that a hierarchical tree with less (or occasionally more) structure than the original better fit the data and so could be preferred for comparative analyses.",
    url = "https://doi.org/10.1111/j.0014-3820.2003.tb00285.x",
    doi = "10.1111/j.0014-3820.2003.tb00285.x",
    openalex = "W1990221500",
    references = "doi1010079781489945419, doi1010160164070486900273, doi10103844766, doi101086284325, doi101086286013, doi101086303327, doi101086343873, doi101086physzool67430163866, doi101093auk1002507, doi101093oso97801985464120010001, doi101093sysbio41118, doi101098rstb19890106, doi101111j001438202000tb00026x, doi101111j001438202001tb00731x, doi1023072412182, doi102307jctt1xp3v3r, doi104159harvard9780674865327, doi105860choice295104, doi105860choice304983, openalexw1550375751, openalexw1558456135"
}

@article{doi101086422893,
    author = "Sih, Andrew and Bell, Alison M. and Johnson, J. Chadwick and Ziemba, Robert",
    title = "Behavioral Syndromes: An Integrative Overview",
    year = "2004",
    journal = "The Quarterly Review of Biology",
    abstract = "A behavioral syndrome is a suite of correlated behaviors expressed either within a given behavioral context (e.g., correlations between foraging behaviors in different habitats) or across different contexts (e.g., correlations among feeding, antipredator, mating, aggressive, and dispersal behaviors). For example, some individuals (and genotypes) might be generally more aggressive, more active or bold, while others are generally less aggressive, active or bold. This phenomenon has been studied in detail in humans, some primates, laboratory rodents, and some domesticated animals, but has rarely been studied in other organisms, and rarely examined from an evolutionary or ecological perspective. Here, we present an integrative overview on the potential importance of behavioral syndromes in evolution and ecology. A central idea is that behavioral correlations generate tradeoffs; for example, an aggressive genotype might do well in situations where high aggression is favored, but might be inappropriately aggressive in situations where low aggression is favored (and vice versa for a low aggression genotype). Behavioral syndromes can thereby result in maladaptive behavior in some contexts, and potentially maintain individual variation in behavior in a variable environment. We suggest terminology and methods for studying behavioral syndromes, review examples, discuss evolutionary and proximate approaches for understanding behavioral syndromes, note insights from human personality research, and outline some potentially important ecological implications. Overall, we suggest that behavioral syndromes could play a useful role as an integrative bridge between genetics, experience, neuroendocrine mechanisms, evolution, and ecology.",
    url = "https://doi.org/10.1086/422893",
    doi = "10.1086/422893",
    openalex = "W2097789622",
    references = "doi101016s0169534701021012, doi101023a1012294324713, doi101093oso97801985774160010001, doi101111j155856461985tb00391x, doi101126science21044731041, doi1018901051076120000100689bicegc20co2"
}

@article{doi101111j14610248200400608x,
    author = "Leibold, Mathew A. and Holyoak, Marcel and Mouquet, Nicolas and Amarasekare, Priyanga and Chase, Jonathan M. and Hoopes, Martha F. and Holt, Robert D. and Shurin, Jonathan B. and Law, Richard and Tilman, David and Loreau, Michel and Gonzalez, Andrew",
    title = "The metacommunity concept: a framework for multi‐scale community ecology",
    year = "2004",
    journal = "Ecology Letters",
    abstract = "Abstract The metacommunity concept is an important way to think about linkages between different spatial scales in ecology. Here we review current understanding about this concept. We first investigate issues related to its definition as a set of local communities that are linked by dispersal of multiple potentially interacting species. We then identify four paradigms for metacommunities: the patch‐dynamic view, the species‐sorting view, the mass effects view and the neutral view, that each emphasizes different processes of potential importance in metacommunities. These have somewhat distinct intellectual histories and we discuss elements related to their potential future synthesis. We then use this framework to discuss why the concept is useful in modifying existing ecological thinking and illustrate this with a number of both theoretical and empirical examples. As ecologists strive to understand increasingly complex mechanisms and strive to work across multiple scales of spatio‐temporal organization, concepts like the metacommunity can provide important insights that frequently contrast with those that would be obtained with more conventional approaches based on local communities alone.",
    url = "https://doi.org/10.1111/j.1461-0248.2004.00608.x",
    doi = "10.1111/j.1461-0248.2004.00608.x",
    openalex = "W2129163149",
    references = "doi1010160040580977900429, doi101016s0065250408602883, doi101038260204c0, doi101046j14610248200300530x, doi101086282398, doi101086283817, doi101086284880, doi101093aibsbulletin2214b, doi101093besa153237, doi101093oso97801985406630010001, doi101111j1469185x1977tb01347x, doi101111j20060906759004272x, doi1015159780691206912, doi1015159781400881376, doi1023071439305, doi1023071931600, doi1023071935620, doi1023071939377, doi1023071941447, doi1023072259756, doi1023072389612, doi1023073071998, doi1023074549, doi105860choice332720"
}

@article{doi1016410006356820040540755wateof20co2,
    author = "Ripple, William J. and Beschta, Robert L.",
    title = "Wolves and the Ecology of Fear: Can Predation Risk Structure Ecosystems?",
    year = "2004",
    journal = "BioScience",
    abstract = "Abstract We investigated how large carnivores, herbivores, and plants may be linked to the maintenance of native species biodiversity through trophic cascades. The extirpation of wolves (Canis lupus) from Yellowstone National Park in the mid-1920s and their reintroduction in 1995 provided the opportunity to examine the cascading effects of carnivore–herbivore interactions on woody browse species, as well as ecological responses involving riparian functions, beaver (Castor canadensis) populations, and general food webs. Our results indicate that predation risk may have profound effects on the structure of ecosystems and is an important constituent of native biodiversity. Our conclusions are based on theory involving trophic cascades, predation risk, and optimal foraging; on the research literature; and on our own recent studies in Yellowstone National Park. Additional research is needed to understand how the lethal effects of predation interact with its nonlethal effects to structure ecosystems.",
    url = "https://doi.org/10.1641/0006-3568(2004)054[0755:wateof]2.0.co;2",
    doi = "10.1641/0006-3568(2004)054[0755:wateof]2.0.co;2",
    openalex = "W2174884086",
    references = "doi1016410006356820040540123rconac20co2, doi1018900012965820030841083arotii20co2, doi1023071383287"
}

@article{doi101890040953,
    author = "Fortin, Daniel and Beyer, Hawthorne L. and Boyce, Mark S. and Smith, Douglas W. and Duchesne, Thierry and Mao, Julie S.",
    title = "WOLVES INFLUENCE ELK MOVEMENTS: BEHAVIOR SHAPES A TROPHIC CASCADE IN YELLOWSTONE NATIONAL PARK",
    year = "2005",
    journal = "Ecology",
    abstract = "A trophic cascade recently has been reported among wolves, elk, and aspen on the northern winter range of Yellowstone National Park, Wyoming, USA, but the mechanisms of indirect interactions within this food chain have yet to be established. We investigated whether the observed trophic cascade might have a behavioral basis by exploring environmental factors influencing the movements of 13 female elk equipped with GPS radio collars. We developed a simple statistical approach that can unveil the concurrent influence of several environmental features on animal movements. Paths of elk traveling on their winter range were broken down into steps, which correspond to the straight-line segment between successive locations at 5-hour intervals. Each observed step was paired with 200 random steps having the same starting point, but differing in length and/or direction. Comparisons between the characteristics of observed and random steps using conditional logistic regression were used to model environmental features influencing movement patterns. We found that elk movements were influenced by multiple factors, such as the distance from roads, the presence of a steep slope along the step, and the cover type in which they ended. The influence of cover type on elk movements depended on the spatial distribution of wolves across the northern winter range of the park. In low wolf-use areas, the relative preference for end point locations of steps followed: aspen stands > open areas > conifer forests. As the risks of wolf encounter increased, the preference of elk for aspen stands gradually decreased, and selection became strongest for steps ending in conifer forests in high wolf-use areas. Our study clarifies the behavioral mechanisms involved in the trophic cascade of Yellowstone's wolf-elk-aspen system: elk respond to wolves on their winter range by a shift in habitat selection, which leads to local reductions in the use of aspen by elk.",
    url = "https://doi.org/10.1890/04-0953",
    doi = "10.1890/04-0953",
    openalex = "W2028644412",
    references = "doi101007bf00395696, doi101139z90092, doi1023073801180"
}

@article{doi101666070541,
    author = "Smith, J. Travis and Jackson, Jeremy B. C.",
    title = "Ecology of extreme faunal turnover of tropical American scallops",
    year = "2009",
    journal = "Paleobiology",
    abstract = "The marine faunas of tropical America underwent substantial evolutionary turnover in the past 3 to 4 million years in response to changing environmental conditions associated with the rise of the Isthmus of Panama, but the ecological signature of changes within major clades is still poorly understood. Here we analyze the paleoecology of faunal turnover within the family Pectinidae (scallops) over the past 12 Myr. The fossil record for the southwest Caribbean (SWC) is remarkably complete over this interval. Diversity increased from a low of 12 species ca. 10–9 Ma to a maximum of 38 species between 4 and 3 Ma and then declined to 22 species today. In contrast, there are large gaps in the record from the tropical eastern Pacific (TEP) and diversity remained low throughout the past 10 Myr. Both origination and extinction rates in the SWC peaked between 4 and 3 Ma, and remained high until 2–1 Ma, resulting in a 95\% species level turnover between 3.5 and 2 Ma. The TEP record was too incomplete for meaningful estimates of origination and extinction rates. All living species within the SWC originated within the last 4 Myr, as evidenced by a sudden jump in Lyellian percentages per faunule from nearly zero up to 100\% during this same interval. However, faunules with Lyellian percentages near zero occurred until 1.8 Ma, so that geographic distributions were extraordinarily heterogeneous until final extinction occurred. There were also striking differences in comparative diversity and abundance among major ecological groups of scallops. Free-swimming scallops constituted the most diverse guild throughout most of the last 10 Myr in the SWC, and were always moderately to very abundant. Leptopecten and Argopecten were also highly diverse throughout the late Miocene and early Pliocene, but declined to very few species thereafter. In contrast, byssally attaching scallops gradually increased in both diversity and abundance since their first appearance in our samples from 8–9 Ma and are the most diverse group today. Evolutionary turnover of scallops in the SWC was correlated with strong ecological reorganization of benthic communities that occurred in response to declining productivity and increased development of corals reefs.",
    url = "https://doi.org/10.1666/07054.1",
    doi = "10.1666/07054.1",
    openalex = "W1972912998",
    references = "doi101016s089598119900005x"
}

@article{doi101371journalpone0047819,
    author = "Jorgensen, Salvador J. and Arnoldi, Natalie S. and Estess, Ethan E. and Chapple, Taylor K. and Rückert, Martin and Anderson, Scot D. and Block, Barbara A.",
    title = "Eating or Meeting? Cluster Analysis Reveals Intricacies of White Shark (Carcharodon carcharias) Migration and Offshore Behavior",
    year = "2012",
    journal = "PLoS ONE",
    abstract = "Elucidating how mobile ocean predators utilize the pelagic environment is vital to understanding the dynamics of oceanic species and ecosystems. Pop-up archival transmitting (PAT) tags have emerged as an important tool to describe animal migrations in oceanic environments where direct observation is not feasible. Available PAT tag data, however, are for the most part limited to geographic position, swimming depth and environmental temperature, making effective behavioral observation challenging. However, novel analysis approaches have the potential to extend the interpretive power of these limited observations. Here we developed an approach based on clustering analysis of PAT daily time-at-depth histogram records to distinguish behavioral modes in white sharks (Carcharodon carcharias). We found four dominant and distinctive behavioral clusters matching previously described behavioral patterns, including two distinctive offshore diving modes. Once validated, we mapped behavior mode occurrence in space and time. Our results demonstrate spatial, temporal and sex-based structure in the diving behavior of white sharks in the northeastern Pacific previously unrecognized including behavioral and migratory patterns resembling those of species with lek mating systems. We discuss our findings, in combination with available life history and environmental data, and propose specific testable hypotheses to distinguish between mating and foraging in northeastern Pacific white sharks that can provide a framework for future work. Our methodology can be applied to similar datasets from other species to further define behaviors during unobservable phases.",
    url = "https://doi.org/10.1371/journal.pone.0047819",
    doi = "10.1371/journal.pone.0047819",
    openalex = "W2005177955",
    references = "doi101007bf00005175"
}

@incollection{davey2018behavioral,
    author = "Davey, Graham",
    title = "Behavioral ecology",
    year = "2018",
    booktitle = "Ecological Learning Theory",
    url = "https://doi.org/10.4324/9781315149233-8",
    doi = "10.4324/9781315149233-8",
    pages = "236-254"
}

@incollection{balch2022behavioral,
    author = "Balch, John",
    title = "Behavioral Ecology",
    year = "2022",
    booktitle = "The Routledge Handbook of Evolutionary Approaches to Religion",
    url = "https://doi.org/10.4324/b23047-27",
    doi = "10.4324/b23047-27",
    pages = "362-381"
}

@article{ohanlon2025behavior,
    author = "O’Hanlon, James C and Khan, M Kawsar and Griffith, Simon C and Chown, Steven L and Cooper, Christine Elizabeth and Duursma, Daisy Englert and Gallagher, Rachael V and Sgrò, Carla M and While, Geoff M and Herberstein, Marie E",
    title = "Behavior across time and space – how large scale “trait-based” approaches can shape behavioral ecology",
    year = "2025",
    journal = "Behavioral Ecology",
    abstract = "Understanding the responses of organisms to environmental change is critical to tackling the grand challenges of 21st century biology. Fields such as ecophysiology and ecology have embraced these challenges and “re-invented” themselves in part by shifting the scale of scientific enquiry and utilizing large-scale comparative approaches. Behavioural research has not yet realized this potential to the same extent. In this paper, we argue that adopting a trait-based approach at large spatial, temporal and taxonomic scales can advance the field of behavioral ecology and address emerging questions in biology. We surveyed the literature in relevant ecology and behavior journals between 1981 and 2020 and found that ecological journals have changed markedly over time, specifically in their focus on understanding interspecific trait variation at broad taxonomic, spatial and temporal scales. This pattern is not apparent for animal behavior, where intra-specific and often intra-population scale of scientific enquiry has mostly been the focus over the last four decades. We argue that behavioral plasticity can be a critical first response to environmental change that might buffer or even lower the risk of extinction. To estimate the capacity of populations or species to respond to change behaviorally, we propose a comparative approach- spatially, temporally or taxonomically- that systematically captures variation in key traits with broad implications for conservation and community ecology. Further, we provide guidance in the methods and resources required to apply a trait-based approach to animal behavior.",
    url = "https://doi.org/10.1093/beheco/araf073",
    doi = "10.1093/beheco/araf073",
    number = "4",
    openalex = "W4411777842",
    volume = "36",
    references = "doi1010079780387874586, doi101016jbiocon201901020, doi101038nature02403, doi101038nature09670, doi101111j001438202003tb00285x, doi101111j14610248200901285x, doi101111j20070906759005171x, doi101126science1251817, doi101371journalpone0185809, doi1023071939924, ohanlon2025behavior"
}

@article{doi101093behecoarag025,
    author = "Guillemin, Liselotte and Sandoz, J. C. and Rödel, Heiko G and d’Ettorre, Patrizia",
    title = "Individual differences in behavior: a multi-trait study comparing different ant species",
    year = "2026",
    journal = "Behavioral Ecology",
    abstract = "Abstract Animal personality, characterized by consistent individual variation in behavioral expression, and behavioral syndromes, underlying correlations between behaviors, have been studied in numerous vertebrate and invertebrate species, including ants. Yet, research on the evolutionary origins of this (co)variation has progressed more slowly than studies showing its occurrence, partly due to varying methodologies preventing comparisons among species. In this study, we applied a comparative framework to investigate patterns of average behaviors and behavioral variation in 4 ant species belonging to 2 different subfamilies: Formica fusca and Camponotus aethiops (Formicinae), Aphaenogaster senilis and Messor barbarus (Myrmicinae). We analyzed 4 behaviors reflecting the responses of ants in foraging or social contexts: exploratory activity, reaction-to-prey, reaction-to-brood, and sociability. All traits except sociability showed moderate to high repeatability in each species. When comparing species for each personality trait, in terms of both average behavioral expression and among-individual variance, we found evidence that phylogeny possibly influences these parameters in the reaction-to-prey test, with closely related species showing more similar behavioral profiles. Other differences among species appear to stem from ecological differences. Robust evidence of behavioral syndromes was found only in A. senilis, with clear correlation between reaction-to-brood and reaction-to-prey, as well as between reaction-to-brood and exploratory activity. Taken together, our findings suggest that our protocol is promising for assessing personality traits across different ant species and that phylogeny may influence the expression of inter-individual differences in certain behavioral traits. These results call for further studies exploring the evolutionary basis of animal personality within a comparative framework.",
    url = "https://doi.org/10.1093/beheco/arag025",
    doi = "10.1093/beheco/arag025",
    openalex = "W7131359716",
    references = "ohanlon2025behavior"
}

@misc{crossrefNonebehavioral,
    title = "Behavioral ecology",
    year = "None",
    booktitle = "AccessScience",
    url = "https://doi.org/10.1036/1097-8542.077250",
    doi = "10.1036/1097-8542.077250"
}