Animal Group Size Research Articles

Do you see what I see? Variation in detection, identification and enumeration of mammals during transect surveys

AbstractEffective monitoring, management and conservation of wildlife axiomatically depend on accurate data but causes of variation, including inter‐observer variation, are rarely explicitly quantified. Here, under controlled conditions, we demonstrate considerable variation in detection, identification and enumeration of (theoretically) readily identifiable African mammals at a reserve with a known species assemblage. Detection: 97.8% of sightings were missed by ≥1 observer; frequency of detection was affected by observer ID, detection distance, visibility and animal group size. Identification: just 3/14 species were identified consistently at all sightings. Enumeration: lack of consensus for 60.5% of sightings; consensus likelihood was affected by visibility and group size.

What is simple is actually quite complex: A critical note on terminology in the domain of language and communication.

On the surface, the fields of animal communication and human linguistics have arrived at conflicting theories and conclusions with respect to the effect of social complexity on communicative complexity. For example, an increase in group size is argued to have opposite consequences on human versus animal communication systems: although an increase in human community size leads to some types of language simplification, an increase in animal group size leads to an increase in signal complexity. But do human and animal communication systems really show such a fundamental discrepancy? Our key message is that the tension between these two adjacent fields is the result of (a) a focus on different levels of analysis (namely, signal variation or grammar-like rules) and (b) an inconsistent use of terminology (namely, the terms "simple" and "complex"). By disentangling and clarifying these terms with respect to different measures of communicative complexity, we show that although animal and human communication systems indeed show some contradictory effects with respect to signal variability, they actually display essentially the same patterns with respect to grammar-like structure. This is despite the fact that the definitions of complexity and simplicity are actually aligned for signal variability, but diverge for grammatical structure. We conclude by advocating for the use of more objective and descriptive terms instead of terms such as "complexity," which can be applied uniformly for human and animal communication systems-leading to comparable descriptions of findings across species and promoting a more productive dialogue between fields. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Dynamics of periodically forced finite N-oscillators, with implications for the social synchronization of animal rest-activity rhythms.

The possible mechanisms for the synchronization of rest-activity rhythms of individual animals living in groups is a relatively understudied question; synchronized rhythms could occur by entrainment of individuals to a common external force and/or by social synchronization between individuals. To gain insight into this question, we explored the synchronization dynamics of populations of globally coupled Kuramoto oscillators and analyzed the effects of a finite oscillator number (N) and the variable strengths of their periodic forcing (F) and mutual coupling (K). We found that increasing N promotes entrainment to a decreasing value of F, but that F could not be reduced below a certain level determined by the number of oscillators and the distribution width of their intrinsic frequencies. Our analysis prompts some simple predictions of ecologically optimal animal group sizes under differing natural conditions.

Variability in the estimation of ungulate group sizes complicates ecological inference.

Foundational work has examined adaptive social behavior in animals in relation to the costs and benefits of group living. Within this context, a “group” of animals represents an organizational unit that is integral to the study of animal ecology and evolution.Definitions of animal group sizes are often subjective with considerable variability within and across species. However, investigations of both the extent and implications of such variability in the estimation of animal group sizes are currently lacking.Selecting ungulates as a case study, we conducted a literature review to assess prevailing practices used to determine group sizes among terrestrial Cetartiodactyla and Perissodactyla. Via this process, we examined group size definitions for 61 species across 171 peer‐reviewed studies published between 1962 and 2018.These studies quantified group sizes via estimation of ungulate aggregations in space and time. Spatial estimates included a nearest neighbor distance ranging from 1.4 m to 1,000 m, and this variation was partially explained by a weak positive correlation (|r| = .4, p < .003) with the body size of the ungulate research subjects. The temporal extent over which group size was estimated was even broader, ranging from three minutes to 24 hr.The considerable variability in ungulate group size estimation that we observed complicates efforts to not only compare and replicate studies but also to evaluate underlying theories of group living. We recommend that researchers: (a) clearly describe the spatiotemporal extents over which they define ungulate group sizes, (b) highlight foundational empirical and ecological rationale for these extents, and (c) seek to align such extents among individual species so as to facilitate cross‐system comparisons of ungulate group size dynamics. We believe an integrative approach to ungulate group size estimation would readily facilitate replication, comparability, and evaluation of competing hypotheses examining the tradeoffs of animal sociality.

Drawn out of the shadows: Surveying secretive forest species with camera trap distance sampling

Abstract With animal species disappearing at unprecedented rates, we need an efficient monitoring method providing reliable estimates of population density and abundance, critical for the assessment of population status and trend. We deployed 160 camera traps (CTs) systematically over 743 locations covering 17,127 km2 of evergreen lowland rainforest of Salonga National Park, block South, Democratic Republic of the Congo. We evaluated the applicability of CT distance sampling (CTDS) to species different in size and behaviour. To improve precision of estimates, we evaluated two methods estimating species' availability (‘A’) for detection by CTs. We recorded 16,700 video clips, revealing 43 different animal taxa. We estimated densities of 14 species differing in physical, behavioural and ecological traits, and extracted species‐specific availability from available video footage using two methods (a) ‘ACa’ (Cappelle et al. [2019] Am. J. Primatol., 81, e22962) and (b) ‘ARo’ (Rowcliffe et al. [2014] Methods Ecol. Evol. 5, 1170). With sample sizes being large enough, we found minor differences between ACa and ARo in estimated densities. In contrast, low detectability and reactivity to the camera were main sources of bias. CTDS proved efficient for estimating density of homogenously rather than patchily distributed species. Synthesis and applications. Our application of camera trap distance sampling (CTDS) to a diverse vertebrate community demonstrates the enormous potential of this methodology for surveys of terrestrial wildlife, allowing rapid assessments of species' status and trends that can translate into effective conservation strategies. By providing the first estimates of understudied species such as the Congo peafowl, the giant ground pangolin and the cusimanses, CTDS may be used as a tool to revise these species' conservation status in the IUCN Red List of Threatened Species. Based on the constraints we encountered, we identify improvements to the current application, enhancing the general applicability of this method.

A Markov jump process modelling animal group size statistics

We translate a coagulation-framentation model, describing the dynamics of animal group size distributions, into a model for the population distribution and associate the \blue{nonlinear} evolution equation with a Markov jump process of a type introduced in classic work of H.~McKean. In particular this formalizes a model suggested by H.-S. Niwa [J.~Theo.~Biol.~224 (2003)] with simple coagulation and fragmentation rates. Based on the jump process, we develop a numerical scheme that allows us to approximate the equilibrium for the Niwa model, validated by comparison to analytical results by Degond et al. [J.~Nonlinear Sci.~27 (2017)], and study the population and size distributions for more complicated rates. Furthermore, the simulations are used to describe statistical properties of the underlying jump process. We additionally discuss the relation of the jump process to models expressed in stochastic differential equations and demonstrate that such a connection is justified in the case of nearest-neighbour interactions, as opposed to global interactions as in the Niwa model.

Self-similar Spreading in a Merging-Splitting Model of Animal Group Size

In a recent study of certain merging-splitting models of animal-group size (Degond et al., J. Nonl. Sci. 27 (2017) 379), it was shown that an initial size distribution with infinite first moment leads to convergence to zero in weak sense, corresponding to unbounded growth of group size. In the present paper we show that for any such initial distribution with a power-law tail, the solution approaches a self-similar spreading form. A one-parameter family of such self-similar solutions exists, with densities that are completely monotone, having power-law behavior in both small and large size regimes, with different exponents.

Numerical approximation of a coagulation-fragmentation model for animal group size statistics

We study numerically a coagulation-fragmentation model derived by Niwa [17] and further elaborated by Degond et al. [5]. In [5] a unique equilibrium distribution of group sizes is shown to exist in both cases of continuous and discrete group size distributions. We provide a numerical investigation of these equilibria using three different methods to approximate the equilibrium: a recursive algorithm based on the work of Ma et. al. [12], a Newton method and the resolution of the time-dependent problem. All three schemes are validated by showing that they approximate the predicted small and large size asymptotic behaviour of the equilibrium accurately. The recursive algorithm is used to investigate the transition from discrete to continuous size distributions and the time evolution scheme is exploited to show uniform convergence to equilibrium in time and to determine convergence rates.

Comparing radio-tracking and visual detection methods to quantify group size measures

Abstract1. Average values of animal group sizes are prone to be overestimated in traditional field studies because small groups and singletons are easier to overlook than large ones. This kind of bias also applies for the method of locating groups by tracking previously radio-collared individuals in the wild. If the researcher randomly chooses a collared animal to locate a group to visit, a large group has higher probability to be selected than a small one, simply because it has more members.2. The question arises whether location of groups by means of finding collared animals has smaller or greater bias than searching for groups by visual observation. If the bias is smaller or same, this method can be recommended for finding groups. However, such a comparison cannot be made by speculation, only by empirical investigation.3. The present study compares the two methods empirically, by statistically comparing group size measures (mean, median, quantiles, frequency distribution, and ‘typical group size’) between two data sets. These data sets comprise of Rocky Mountain mule deer group size values collected in the same area during the same period of time, referring either to groups located by the traditional ‘search and observe method’ or located by tracking formerly collared individuals.4. All group size measures are statistically similar in the two samples, thus we conclude that the two methods yielded similar biases. Although the true group size measures are not known, we presume that both methods have overestimated them. We propose that these results do not necessary apply to other species, thus cannot be generalized. The reason for this is that bias may depend on factors specific to the species: bias of visual observation may depend on how well the species conceals itself in the existing habitat, and the bias associated with finding groups using collared animals is likely dependent on group size distribution and also on the proportion of collared animals in the population.

Managing more than the mean: using quantile regression to identify factors related to large elk groups

Summary Animal group size distributions are often right‐skewed, whereby most groups are small, but most individuals occur in larger groups that may also disproportionately affect ecology and policy. In this case, examining covariates associated with upper quantiles of the group size distribution could facilitate better understanding and management of large animal groups.We studied wintering elk groups in Wyoming, where group sizes span several orders of magnitude, and issues of disease, predation and property damage are affected by larger group sizes. We used quantile regression to evaluate relationships between the group size distribution and variables of land use, habitat, elk density and wolf abundance to identify conditions important to larger elk groups.We recorded 1263 groups ranging from 1 to 1952 elk and found that across all quantiles of group size, group sizes were larger in open habitat and on private land, but the largest effect occurred between irrigated and non‐irrigated land [e.g. the 90th quantile group size increased by 135 elk (95% CI = 42, 227) on irrigation].Only upper quantile group sizes were positively related to broad‐scale measures of elk density and wolf abundance. For wolf abundance, this effect was greater on elk groups found in open habitats and private land than those in closed habitats or public land. If we had limited our analysis to mean or median group sizes, we would not have detected these effects. Synthesis and applications. Our analysis of elk group size distributions using quantile regression suggests that private land, irrigation, open habitat, elk density and wolf abundance can affect large elk group sizes. Thus, to manage larger groups by removal or dispersal of individuals, we recommend incentivizing hunting on private land (particularly if irrigated) during the regular and late hunting seasons, promoting tolerance of wolves on private land (if elk aggregate in these areas to avoid wolves) and creating more winter range and varied habitats. Relationships to the variables of interest also differed by quantile, highlighting the importance of using quantile regression to examine response variables more completely to uncover relationships important to conservation and management.

Incorporating detection probability into northern Great Plains pronghorn population estimates

ABSTRACTPronghorn (Antilocapra americana) abundances commonly are estimated using fixed‐wing surveys, but these estimates are likely to be negatively biased because of violations of key assumptions underpinning line‐transect methodology. Reducing bias and improving precision of abundance estimates through use of detection probability and mark‐resight models may allow for more responsive pronghorn management actions. Given their potential application in population estimation, we evaluated detection probability and mark‐resight models for use in estimating pronghorn population abundance. We used logistic regression to quantify probabilities that detecting pronghorn might be influenced by group size, animal activity, percent vegetation, cover type, and topography. We estimated pronghorn population size by study area and year using mixed logit‐normal mark‐resight (MLNM) models. Pronghorn detection probability increased with group size, animal activity, and percent vegetation; overall detection probability was 0.639 (95% CI = 0.612–0.667) with 396 of 620 pronghorn groups detected. Despite model selection uncertainty, the best detection probability models were 44% (range = 8–79%) and 180% (range = 139–217%) greater than traditional pronghorn population estimates. Similarly, the best MLNM models were 28% (range = 3–58%) and 147% (range = 124–180%) greater than traditional population estimates. Detection probability of pronghorn was not constant but depended on both intrinsic and extrinsic factors. When pronghorn detection probability is a function of animal group size, animal activity, landscape complexity, and percent vegetation, traditional aerial survey techniques will result in biased pronghorn abundance estimates. Standardizing survey conditions, increasing resighting occasions, or accounting for variation in individual heterogeneity in mark‐resight models will increase the accuracy and precision of pronghorn population estimates. © 2013 The Wildlife Society.

Monitoring an Endangered savannah ungulate, Grevy's zebraEquus grevyi: choosing a method for estimating population densities

AbstractMethods that accurately estimate animal abundance or density are crucial for wildlife management. Although numerous techniques are available, there have been few comparisons of the precision and cost-effectiveness of different approaches. We assess the precision and cost of three methods for estimating densities of the Endangered Grevy's zebraEquus grevyi. We compare distance sampling and photographic capture–recapture, and a new technique, the random encounter model (REM) that uses camera-trap encounter rates to estimate density. All three methods provide comparable density estimates for Grevy's zebra and are preferable to the common practice of raw counts. Photographic capture–recapture is the most precise and line-transect distance sampling the least precise. Line transects and photographic capture–recapture surveys are cost-effective in the first year and REM is most cost-effective in the long-term. The methods used here for Grevy's zebra may be applied to other rangeland ungulates. We suggest that for single species monitoring programmes in which individuals can be identified, photographic capture–recapture surveys may be the preferred method for estimating wildlife abundances. When encounter rates are low, distance sampling lacks the precision of the other methods but its cost advantage may make it appropriate for long-term or multi-species monitoring programmes. The REM is an efficient and precise method of estimating densities but has high initial equipment costs. We believe REM has the potential to work well for many species but it requires independent estimates of animal movements and group size.

Positive impact of pre-stroke surgery on survival following transient focal ischemia in hypertensive rats

We describe a positive influence of pre-stroke surgery on recovery and survival in a commonly used experimental stroke model. Two groups of male, stroke-prone spontaneously hypertensive rats (SHRSPs) underwent transient middle cerebral artery occlusion (tMCAO). Group 1 underwent the procedure without any prior intervention whilst group 2 had an additional general anaesthetic 6 days prior to tMCAO for a cranial burrhole and durotomy. Post-stroke recovery was assessed using a 32 point neurological deficit score and tapered beam walk and infarct volume determined from haematoxylin–eosin stained sections. In group 2 survival was 92% (n=12) versus 67% in group 1 (n=18). In addition, post-tMCAO associated weight loss was significantly reduced in group 2. There was no significant difference between the two groups in experimental outcomes: infarct volume (Group 1 317±18.6mm3 versus Group 2 332±20.4mm3), and serial (day 0–14 post-tMCAO) neurological deficit scores and tapered-beam walk test. Drilling a cranial burrhole under general anaesthesia prior to tMCAO in SHRSP reduced mortality and gave rise to infarct volumes and neurological deficits similar to those recorded in surviving Group 1 animals. This methodological refinement has significant implications for animal welfare and group sizes required for intervention studies.

Effect of genetic homogeneity on behavioural variability in an object recognition test in cloned Göttingen minipigs

The number of animals used in research should be limited as much as possible. Among cloned animals, genetic variation is minimal and to the extent that behaviour is genetically determined inter-individual variability is expected to be higher among naturally bred animals. However, the cloning procedure per se might affect the resultant phenotype leading to phenotypic variations independent of the genetic background. Recently, cloned Göttingen minipigs carrying a mutation for Alzheimer's disease have been produced. In order to document the development of Alzheimer's disease symptoms, these pigs were subjected to a behavioural test of memory, the spontaneous object recognition test, from an early age. At ages 1 and 2 years no evidence of memory decline was found, yet the data showed striking behavioural variability among the cloned groups. The aim of the present study was to investigate effects of genetic homogeneity on variability of cloned minipigs compared with non-cloned controls regarding behavioural variables in a cognitive test, namely the spontaneous object recognition test.Significant differences in the variability between the cloned and control pigs were found in five out of 24 behavioural variables. The clones showed lower variability for four of the measures, whereas the variability was increased for one variable. No significant difference between the cloned and the control group was found for the remaining 19 variables (79%). In 14 of these, the standard deviation was numerically greater for the control group compared to the cloned group, indicating that variation may be less in cloned animals, but not demonstrable with the small group size of the present study (n=6 for each of the two groups tested).Overall, this study failed to show unambiguously that variability in these behavioural variables in cloned minipigs is less than in naturally bred control subjects and therefore does not directly support the hypothesis that cloning may be used to replicate animals in order to reduce group size in experimentation animals.

Cone opsins and response of female chamois (Rupicapra rupicapra) to differently coloured raincoats

Alpine species are often exposed to intense levels of human recreational activities. Exactly how human disturbances influence the behaviour of these species is still open to much debate. For example, little is known regarding how the colourful clothing often worn by tourists influences the behaviour of animals. Tourists wearing colourful clothing may be more conspicuous to local wildlife and thus cause more disturbances. We therefore investigated this question in female chamois (Rupicapra rupicapra) in the Swiss Alps. We firstly investigated, via a morphological and an immunohistochemical approach, whether chamois are likely to have colour vision and would therefore be more likely to respond to different coloured clothing. We detected evidence of two cone types—short-wavelength-sensitive cones (S-cones, JH 455) and middle-wavelength-sensitive cones (M-cones, JH492) in the chamois retina—suggesting that chamois have dichromatic vision, similar to other ungulates. Secondly, via behavioural assays where a person wearing one of three coloured coats commonly worn by tourists (red, yellow and blue) approached a female chamois, we show that neither the alert and flight initiation distance nor the site of refuge were influenced by the raincoat colour. In addition, behavioural responses of the chamois were neither influenced by animal group size nor the presence of kids nor the time of the experiment. The results suggest that, although chamois possess colour vision, they do not react more strongly towards conspicuous colours worn by hikers. We discuss our results in light of what is already known about chamois biology and suggest implications for future studies.

Understanding Animal Group-Size Distributions

One of the most striking aspects of animal groups is their remarkable variation in size, both within and between species. While a number of mechanistic models have been proposed to explain this variation, there are few comprehensive datasets against which these models have been tested. In particular, we only vaguely understand how environmental factors and behavioral activities affect group-size distributions. Here we use observations of House sparrows (Passer domesticus) to investigate the factors determining group-size distribution. Over a wide range of conditions, we observed that animal group sizes followed a single parameter distribution known as the logarithmic distribution. This single parameter is the mean group size experienced by a randomly chosen individual (including the individual itself). For sparrows, the experienced mean group size, and hence the distribution, was affected by four factors: morning temperature, place, behavior and the degree of food spillage. Our results further indicate that the sparrows regulate the mean group size they experience, either by groups splitting more or merging less when local densities are high. We suggest that the mean experienced group size provides a simple but general tool for assessing the ecology and evolution of grouping.

An inter-laboratory retrospective analysis of immunotoxicological endpoints in non-human primates: T-cell-dependent antibody responses

The Immunotoxicology Technical Committee of HESI sponsored a retrospective analysis of T-cell-dependent antibody responses in non-human primates (NHP). Antibody responses to keyhole limpet hemocyanin (KLH), tetanus toxoid (TT), and/or sheep red blood cells (SRBC) in 178 NHP (from 8 sponsors, 13 testing sites, 30 studies) were statistically analyzed. Rates of positive or negative anti-KLH, -TT, and -SRBC primary and secondary IgM and IgG responses were compared. The influence of gender, country of origin, and previous immunization with a different antigen on response rate and kinetics of anti-KLH and anti-TT responses were analyzed. In addition, the magnitude of the antibody responses and the impact of the above-mentioned factors were analyzed. In addition, based upon the inter-individual variability of the peak response values, power calculations were conducted. The analysis demonstrated that the rates of positive responses were similar between the two genders, were high for KLH, SRBC, and TT challenges by 21 days following immunization (87, 100, and 84%, respectively, for IgGs) and did not include statistically significant differences based on NHP country of origin. Mean peak secondary responses were greater than peak primary responses; the magnitude of the response to KLH was increased by incomplete Freund’s adjuvant (IFA). Gender had little effect on the magnitude and variability of these responses. KLH and TT were associated with similar inter-animal variability, whereas in some situations KLH responses were less variable than responses to SRBC. The data suggested that inter-animal variability with KLH was similar with or without IFA. Power analysis illustrated that animal group sizes of typical standard toxicology studies (generally ≤ 4/sex) are likely to detect only fairly large treatment effects. However, combining males and females, when appropriate, will improve the power: an N of 8 to 12 could detect ≤ 3.1-fold differences in anti-KLH IgG responses.

A first principles derivation of animal group size distributions

Several empirical studies have shown that the animal group size distribution of many species can be well fit by power laws with exponential truncation. A striking empirical result due to Niwa is that the exponent in these power laws is one and the truncation is determined by the average group size experienced by an individual. This distribution is known as the logarithmic distribution. In this paper we provide first principles derivation of the logarithmic distribution and other truncated power laws using a site-based merge and split framework. In particular, we investigate two such models. Firstly, we look at a model in which groups merge whenever they meet but split with a constant probability per time step. This generates a distribution similar, but not identical to the logarithmic distribution. Secondly, we propose a model, based on preferential attachment, that produces the logarithmic distribution exactly. Our derivation helps explain why logarithmic distributions are so widely observed in nature. The derivation also allows us to link splitting and joining behavior to the exponent and truncation parameters in power laws.

Societal concern related to stocking density, pen size and group size in farm animal production

Farm animal welfare has become increasingly important and relevant from the societal point of view throughout the livestock production chain, with space allowance for farm animals being one of its most topical and vividly debated sub-matters. Anticipating on the increasing power of the citizens' voice in Western society, this study focuses on societal concerns related to space allowance in Flemish animal production. This paper reports a quantitative analysis of cross-sectional surveys conducted among citizens of Flanders during 2000–2002 ( n = 521) and during 2006 ( n = 459). Compared with other animal welfare aspects, the survey participants perceived stocking densities and pen sizes as very essential for attaining an acceptable level of farm animal welfare. In addition, they evaluated stocking densities and pen sizes in current Flemish animal production as inadequate. Combined these results indicate that societal concern, defined as the difference between perceived importance and evaluative belief, is very high for stocking density and pen size, though less for group size. Societal concern about stocking density was found to be higher among younger people and households without children, while societal concern was not significantly associated with living environment (rural versus urban). Gender (higher concern among women in dataset 2006) and educational (higher concern among higher educated people in dataset 2000–2002) differences were not consistently significant in all datasets. The findings from this study suggest that larger gains in terms of animal welfare image among the public can be realised from providing farm animals with more space relative to reducing animal group size.

Estimating animal density using camera traps without the need for individual recognition

Summary 1Density estimation is of fundamental importance in wildlife management. The use of camera traps to estimate animal density has so far been restricted to capture–recapture analysis of species with individually identifiable markings. This study developed a method that eliminates the requirement for individual recognition of animals by modelling the underlying process of contact between animals and cameras. 2The model provides a factor that linearly scales trapping rate with density, depending on two key biological variables (average animal group size and day range) and two characteristics of the camera sensor (distance and angle within which it detects animals). 3We tested the approach in an enclosed animal park with known abundances of four species, obtaining accurate estimates in three out of four cases. Inaccuracy in the fourth species was because of biased placement of cameras with respect to the distribution of this species. 4Synthesis and applications. Subject to unbiased camera placement and accurate measurement of model parameters, this method opens the possibility of reduced labour costs for estimating wildlife density and may make estimation possible where it has not been previously. We provide guidelines on the trapping effort required to obtain reasonably precise estimates.