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Daniel Rubenstein
  • Princeton, New Jersey
Research Interests:
Though morphologically very similar, equids across the extant species occupy ecological niches that are surprisingly non-overlapping. Occupancy of these distinct niches appears related to subtle physiological and behavioural adaptations... more
Though morphologically very similar, equids across the extant species occupy ecological niches that are surprisingly non-overlapping. Occupancy of these distinct niches appears related to subtle physiological and behavioural adaptations which, in turn, correspond to significant differences in the social behaviours and emergent social systems characterizing the different species. Although instances of intraspecific behavioural variation in equids demonstrate that the same body plan can support a range of social structures, each of these morphologically similar species generally shows robust fidelity to its evolved social system. The pattern suggests a subtle relationship between physiological phenotypes and behavioural flexibility. While environmental conditions can vary widely within relatively short temporal or spatial scales, physiological changes and changes to the behaviours that regulate physiological processes, are constrained to longer cycles of adaptation. Physiology is then...
Here, we revise Pietraszewski's model of groups by assigning participant pairs with two triplets, denoting: (1) the type of game that models the interaction, (2) its critical switching point between alternatives (i.e., the game's... more
Here, we revise Pietraszewski's model of groups by assigning participant pairs with two triplets, denoting: (1) the type of game that models the interaction, (2) its critical switching point between alternatives (i.e., the game's similarity threshold), and (3) the perception of strategic similarity with the opponent. These triplets provide a set of primitives that accounts for individuals' strategic motivations and observed behaviors.
conservation status Scientific name: Equus burchellii Gray Equus burchellii boehmi
Individual recognition can be facilitated by creating representations of familiar individuals, whereby information from signals in multiple sensory modalities become linked. Many vertebrate species use auditory–visual matching to... more
Individual recognition can be facilitated by creating representations of familiar individuals, whereby information from signals in multiple sensory modalities become linked. Many vertebrate species use auditory–visual matching to recognize familiar conspecifics and heterospecifics, but we currently do not know whether representations of familiar individuals incorporate information from other modalities. Ring-tailed lemurs ( Lemur catta ) are highly visual, but also communicate via scents and vocalizations. To investigate the role of olfactory signals in multisensory recognition, we tested whether lemurs can recognize familiar individuals through matching scents and vocalizations. We presented lemurs with female scents that were paired with the contact call either of the female whose scent was presented or of another familiar female from the same social group. When the scent and the vocalization came from the same individual versus from different individuals, females showed greater i...
Page 1. 10 D. RUBENSTEIN, R. BARNETT, R. RIDGELY & P. KLOPFER IBIS 119 ADAPTIVE ADVANTAGES OF MIXED-SPECIES FEEDING FLOCKS AMONG SEED-EATING FINCHES IN COSTA RICA DANIEL I. RUBENSTEIN ...
In this study, we investigated the role of assessment time in group size discrimination and in particular the trade-off between the time cost involved in gathering information and the potential benefits derived from the acquired... more
In this study, we investigated the role of assessment time in group size discrimination and in particular the trade-off between the time cost involved in gathering information and the potential benefits derived from the acquired information. In a first experiment, we presented individual chub, Semotilus atromaculatus, with a choice between 4 vs 4 and 8 vs 8 conspecific stimulus fish. After release we recorded the time taken by test fish to make a choice between the two stimulus shoals in the presence and absence of a fright stimulus. Test fish significantly reduced their response time in the presence of a fright stimulus and larger shoals (8 vs 8) were more quickly approached than smaller ones (4 vs 4). In a control experiment, chub were given a choice between an empty cylinder and a shoal (of 4 or 8 fish). By subtracting the response time in the control treatment from that in the choice treatment, we estimated the time test fish spent choosing between stimulus shoals to be 24-55% o...
Although cooperation and trust are essential features for the development of prosperous populations, they also put cooperating individuals at risk for exploitation and abuse. Empirical and theoretical evidence suggests that the solution... more
Although cooperation and trust are essential features for the development of prosperous populations, they also put cooperating individuals at risk for exploitation and abuse. Empirical and theoretical evidence suggests that the solution to the problem resides in the practice of mimicry and imitation, the expectation of opponent’s mimicry and the reliance on similarity indices. Here we fuse the principles of enacted and expected mimicry and condition their application on two similarity indices to produce a model of mimicry and relative similarity. Testing the model in computer simulations of behavioral niches, populated with agents that enact various strategies and learning algorithms, shows how mimicry and relative similarity outperforms all the opponent strategies it was tested against, pushes noncooperative opponents toward extinction, and promotes the development of cooperative populations. The proposed model sheds light on the evolution of cooperation and provides a blueprint fo...
This paper proposes a 5-component detection pipeline for use in a computer vision-based animal recognition system. The end result of our proposed pipeline is a collection of novel annotations of interest (AoI) with species and view-point... more
This paper proposes a 5-component detection pipeline for use in a computer vision-based animal recognition system. The end result of our proposed pipeline is a collection of novel annotations of interest (AoI) with species and view-point labels. These AoIs, for example, could be fed as the focused input data into an appearance-based animal identification system. The goal of our method is to increase the reliability and automation of animal censusing studies and to provide better ecological information to conservationists. Our method is able to achieve a localization mAP of 81.67%, a species and viewpoint annotation classification accuracy of 94.28% and 87.11%, respectively, and an AoI accuracy of 72.75% across 6 animal species of interest. We also introduce the Wildlife Image and Localization Dataset (WILD), which contains 5,784 images and 12,007 labeled annotations across 28 classification species and a variety of challenging, real-world detection scenarios.
Photographs, taken by field scientists, tourists, automated cameras, and incidental photographers, are the most abundant source of data on wildlife today. Wildbook is an autonomous computational system that starts from massive collections... more
Photographs, taken by field scientists, tourists, automated cameras, and incidental photographers, are the most abundant source of data on wildlife today. Wildbook is an autonomous computational system that starts from massive collections of images and, by detecting various species of animals and identifying individuals, combined with sophisticated data management, turns them into high resolution information database, enabling scientific inquiry, conservation, and citizen science. We have built Wildbooks for whales (flukebook.org), sharks (whaleshark.org), two species of zebras (Grevy's and plains), and several others. In January 2016, Wildbook enabled the first ever full species (the endangered Grevy's zebra) census using photographs taken by ordinary citizens in Kenya. The resulting numbers are now the official species census used by IUCN Red List: this http URL In 2016, Wildbook partnered up with WWF to build Wildbook for Sea Turtles, Internet of Turtles (IoT), as well as...
IUCN Red List Category (E. burchellii, E.b. antiquorum, and E.b. boehmi were assessed using version 3.1; all others with version 2.3): Equus burchellii LC Least Concern E. b. boehmi LR Lower Risk E. b. zambesiansis DD Data Deficient (?... more
IUCN Red List Category (E. burchellii, E.b. antiquorum, and E.b. boehmi were assessed using version 3.1; all others with version 2.3): Equus burchellii LC Least Concern E. b. boehmi LR Lower Risk E. b. zambesiansis DD Data Deficient (? extinct in wild) E. b. crawshayi DD Data Deficient (? endangered) E. b. chapmani DD Data Deficient E. b. antiquorum LR Lower Risk E. b. burchellii EX Extinct (1930)
Studies in Behavioural Adaptation Series Editor: John Lazarus, Department of Psychology, The University of Newcastle upon Tyne Gulls and Plovers: The Ecology and Behaviour of Mixed-Species Feeding Groups CJ Barnard and DBA Thompson ...
In several social species, adult associations have been linked to individual fitness. Less is known about offspring associations and the mechanisms by which they may influence fitness. We investigate the effects of sociality on juvenile... more
In several social species, adult associations have been linked to individual fitness. Less is known about offspring associations and the mechanisms by which they may influence fitness. We investigate the effects of sociality on juvenile survival in feral horses (Equus caballus). We use foal degree (number of associates) and foal weight (number of interactions) to assess sociality’s importance to foal survival of a catastrophic event, the gathering and removal of 40% of the horse population. We found that 1) foal degree was a better predictor of foal survival than was foal weight; 2) following the gather, foals with access to at least one parent and some members of their natal groups were more likely to survive than were foals left with no such access; 3) foals with a higher degree both pre- and postgather were more likely to survive; 4) the influence of pregather degree appeared to be more pronounced among foals without access to parent(s) and natal group members; and 5) the influen...
Research Interests:
Research Interests:
Grevy's zebra are endangered species of zebra found in Eastern Africa. Onagers are very common wild Asses widespread from Middle East to India. Both species are equids of roughly the same size, similar physiology, and living in... more
Grevy's zebra are endangered species of zebra found in Eastern Africa. Onagers are very common wild Asses widespread from Middle East to India. Both species are equids of roughly the same size, similar physiology, and living in semiarid environment. Both were previously described as having the same social organization: social fission-fusion species with territorial males and harem females. While traditional social network analysis has revealed some differences in the structure of their interactions, it was not sufficient to fully ...
The time immediately following birth is a period of high predation risk for ungulate neonates. Ungulate mothers exhibit perinatal behaviors that appear to mitigate offspring risk during this time. However, few studies of infant mortality... more
The time immediately following birth is a period of high predation risk for ungulate neonates. Ungulate mothers exhibit perinatal behaviors that appear to mitigate offspring risk during this time. However, few studies of infant mortality include the postpartum period. Therefore, the function and effectiveness of these maternal behaviors are untested. We observed perinatal behavior in 11 Thomson’s gazelle (Eudorcas thomsoni) mother–infant pairs in a free-ranging population under predation pressure. Five of the six fawns that were detected by predators during the perinatal period were killed. Fawn survival therefore depended on avoiding detection by predators. Considered individually, neither prepartum isolation from conspecifics nor birth site selection affected the risk of being detected by a predator. However, analyses revealed two distinct perinatal tactics: mothers either isolated and gave birth in tall grass or remained in their social groups and gave birth in short grass. Both ...
Human health and livelihoods are threatened by declining marine fisheries catches, causing substantial interest in the sources and dynamics of fishing. Catch analyses in individual exclusive economic zones (EEZs) and the high seas are... more
Human health and livelihoods are threatened by declining marine fisheries catches, causing substantial interest in the sources and dynamics of fishing. Catch analyses in individual exclusive economic zones (EEZs) and the high seas are abundant, and research across multiple EEZs is growing. However, no previous studies have systematically compared catches, intranational versus international fish flows, and fishing nations within all of the world’s EEZs and across adjacent and distant EEZs and the high seas to inform “metacoupled” fisheries management. We use the metacoupling framework—a new approach for evaluating human–nature interactions within and across adjacent and distant systems (metacouplings)—to illustrate how fisheries catches were locally, regionally, and globally interconnected in 1950–2014, totaling 5.8 billion metric tons and increasing by 298% (tonnage) and 431% (monetary value) over this time period. Catches by nations in their own EEZs (largest in Peru) and adjacent ...
Following the outbreak of COVID-19 pandemic, governments around the globe coerced their citizens to adhere to preventive health behaviours, aiming to reduce the effective reproduction numbers of the virus. Driven by game theoretic... more
Following the outbreak of COVID-19 pandemic, governments around the globe coerced their citizens to adhere to preventive health behaviours, aiming to reduce the effective reproduction numbers of the virus. Driven by game theoretic considerations and inspired by the work of US National Research Council's Committee on Food Habits (1943) during WWII, and the post-WWII Yale Communication Research Program, the present research shows how to achieve enhanced adherence to health regulations without coercion. To this aim, we combine three elements: (i) indirect measurements, (ii) personalized interventions, and (iii) attitude changing treatments (IMPACT). We find that a cluster of short interventions, such as elaboration on possible consequences, induction of cognitive dissonance, addressing next of kin and similar others and receiving advice following severity judgements, improves individuals' health-preserving attitudes. We propose extending the use of IMPACT under closure periods ...
Group members interact with each other during multiple social behaviours that range from aggressive to affiliative interactions. It is not known, however, whether an individual's suite of social behaviours consistently covaries... more
Group members interact with each other during multiple social behaviours that range from aggressive to affiliative interactions. It is not known, however, whether an individual's suite of social behaviours consistently covaries through time and across different types of social interactions. Consistent social behaviour would be advantageous in groups, especially when individuals need to remember their group members' social roles and preferences in order to keep track of social relationships and predict conspecifics' future behaviour. Here, we address whether social behaviour of ringtailed lemurs, Lemur catta, is consistent through time and across four interaction networks (aggression, grooming, contact calling, scent marking). We quantified variation in social behaviour through four network centrality measures including outdegree, outstrength, betweenness and eigenvector centrality. Comparing lemurs' measures across 2 years revealed that network centrality remained consistent between years. Lemurs' centrality also stayed consistent across interaction networks: individuals with high centrality in one interaction network also had high centrality in the other networks, even when we controlled for sex-based variation in social behaviour. Thus, regardless of their sex, some individuals were highly social and frequently groomed others, initiated aggressive interactions and responded to others' contact calls and scent marks. Lemurs also had preferred social partners they frequently interacted with across years and across multiple behaviours. In particular, lemurs frequently responded to the contact calls and the scent marks of the conspecifics they had frequently groomed. Together, these results demonstrate that individual variation in lemur social behaviour is not context specific, but instead persists through time and across multiple social interactions. Such consistent behaviour provides evidence of social personalities, which may influence individuals' interaction styles, including how socially active they are and with whom they interact.
When predators are removed or suppressed for generations, prey populations tend to increase and when predators are re-introduced, prey densities should fall back to pre-control levels. In cases of apparent competition where there are... more
When predators are removed or suppressed for generations, prey populations tend to increase and when predators are re-introduced, prey densities should fall back to pre-control levels. In cases of apparent competition where there are alternate abundant and rare prey species, rare species may decline further than expected or disappear altogether. Recently, concern about the impact of recovering predator populations on wildlife in Laikipia County, Kenya, has led to questions of whether lions (Panthera leo, IUCN Red List Vulnerable) exert top-down pressure on Grevy's zebra (Equus grevyi, IUCN Red List Endangered). We examined effects of lion predation on Plain's zebra (E. quagga, IUCN Red List Near Threatened) and Grevy's zebra populations in a 2,105 km2 area defined by lion movements. We used line transect surveys to estimate density of Grevy's (0.71/km2) and Plain's (15.9/km2) zebras, and satellite telemetry to measure movements for lions and both zebras. We track...
Understanding how humans and other animals behave in response to changes in their environments is vital for predicting population dynamics and the trajectory of coupled social-ecological systems. Here, we present a novel framework for... more
Understanding how humans and other animals behave in response to changes in their environments is vital for predicting population dynamics and the trajectory of coupled social-ecological systems. Here, we present a novel framework for identifying emergent social behaviours in foragers (including humans engaged in fishing or hunting) in predator-prey contexts based on the exploration difficulty and exploitation potential of a renewable natural resource. A qualitative framework is introduced that predicts when foragers should behave territorially, search collectively, act independently or switch among these states. To validate it, we derived quantitative predictions from two models of different structure: a generic mathematical model, and a lattice-based evolutionary model emphasising exploitation and exclusion costs. These models independently identified that the exploration difficulty and exploitation potential of the natural resource controls the social behaviour of resource exploi...
Strong relationships exist between social connections and information transmission [1-9], where individuals' network position plays a key role in whether or not they acquire novel information [2, 3, 5, 6]. The relationships between... more
Strong relationships exist between social connections and information transmission [1-9], where individuals' network position plays a key role in whether or not they acquire novel information [2, 3, 5, 6]. The relationships between social connections and information acquisition may be bidirectional if learning novel information, in addition to being influenced by it, influences network position. Individuals who acquire information quickly and use it frequently may receive more affiliative behaviors [10, 11] and may thus have a central network position. However, the potential influence of learning on network centrality has not been theoretically or empirically addressed. To bridge this epistemic gap, we investigated whether ring-tailed lemurs' (Lemur catta) centrality in affiliation networks changed after they learned how to solve a novel foraging task. Lemurs who had frequently initiated interactions and approached conspecifics before the learning experiment were more likely...
Effective population size, a central concept in conservation biology, is now routinely estimated from genetic surveys, and can also be theoretically-predicted from demographic, life-history and mating-system hypotheses. However, by... more
Effective population size, a central concept in conservation biology, is now routinely estimated from genetic surveys, and can also be theoretically-predicted from demographic, life-history and mating-system hypotheses. However, by evaluating the consistency of theoretical predictions with empirically-estimated effective size, insights can be gained regarding life-history characteristics, as well as the relative impact of different life-history traits on genetic drift. These insights can be used to design and inform management strategies aimed at increasing effective population size. Here we describe and demonstrate this approach by addressing the conservation of a reintroduced population of Asiatic wild ass (Equus hemionus). We estimate the variance effective size (N) from genetic data (N= 24.3), and we formulate predictions for the impacts on Nof demography, polygyny, female variance in life-time reproductive success, and heritability of female reproductive success. By contrasting...
Animals are predicted to selectively observe and learn from the conspecifics with whom they share social connections. Yet, hardly anything is known about the role of different connections in observation and learning. To address the... more
Animals are predicted to selectively observe and learn from the conspecifics with whom they share social connections. Yet, hardly anything is known about the role of different connections in observation and learning. To address the relationships between social connections, observation and learning, we investigated transmission of information in two raven ( Corvus corax ) groups. First, we quantified social connections in each group by constructing networks on affiliative interactions, aggressive interactions and proximity. We then seeded novel information by training one group member on a novel task and allowing others to observe. In each group, an observation network based on who observed whose task-solving behaviour was strongly correlated with networks based on affiliative interactions and proximity. Ravens with high social centrality (strength, eigenvector, information centrality) in the affiliative interaction network were also central in the observation network, possibly as a ...
When prey are time limited in their access to food, any trade-off involving time should ultimately affect their intake rate. In many herbivores, males and females experience different ecological pressures affecting their survival and... more
When prey are time limited in their access to food, any trade-off involving time should ultimately affect their intake rate. In many herbivores, males and females experience different ecological pressures affecting their survival and reproduction because of differences in morphology, physiology and energy/nutrient requirements. If males and females have different vigilance strategies that affect their intake rates differently, they will suffer different foraging costs. This is particularly relevant in sexually monomorphic herbivores, where the two sexes have similar basal energy/nutrient requirements and risk of predation. We investigated how gender, reproductive status, age, group size, predation risk, and food biomass affected vigilance, intake rate, and their trade-off in a monomorphic species, the plains zebra (Equus quagga). Males were more vigilant than females, and lactating females were less vigilant than other females; the levels of vigilance were low (ca. 10 % of feeding t...
ABSTRACT Sperm competition occurs in the giant water strider. In serial mating experiments using normal and sterilized males, the last male to mate was able to displace, on average, 65% of the sperm of the previous male. Success of second... more
ABSTRACT Sperm competition occurs in the giant water strider. In serial mating experiments using normal and sterilized males, the last male to mate was able to displace, on average, 65% of the sperm of the previous male. Success of second males was, however, highly variable but can be partially explained by relative copulation lengths of the two males. Although there was a tendency for long copulations by the first male to be followed by long copulations by the second, second males could only obtain the ‘last male’ advantage if their copulations were long relative to those of first males. A long latency period occurred between genital contact and the effective transfer of sperm as no viable eggs were produced until copulations lasted at least 15 min. Once sperm transfer began, however, virtually the entire sperm load was delivered instantaneously to the female. Females continued to lay fertile eggs for 24 days after a single copulation, and although this period increased to 30 days after two matings, the difference was not statistically significant. Although the incompleteness of sperm displacement suggests that alternative copulatory strategies among males should be common, the existence of a long latency period before sperm transfer should not only reduce this tendency, but also increase female choosiness.
SUMMARY When creating protected area networks in fragmented landscapes with limited ecological data at hand, one approach is to use design criteria, such as patch size or edge/area ratios, to rank the ecosystem patches' conservation... more
SUMMARY When creating protected area networks in fragmented landscapes with limited ecological data at hand, one approach is to use design criteria, such as patch size or edge/area ratios, to rank the ecosystem patches' conservation value (e.g., habitat integrity, biodiversity) and evaluate protected area network alternatives. Caution is warranted, however, as the relationships between these design criteria and the ecosystem

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