Roger Mundry

Stepwise Model Fitting and Statistical Inference: Turning Noise into Signal Pollution

Statistical inference based on stepwise model selection is applied regularly in ecological, evolutionary, and behavioral research. In addition to fundamental shortcomings with regard to finding the “best” model, stepwise procedures are known to suffer from a multiple‐testing problem, yet the method is still widely used. As an illustration of this problem, we present results of a simulation study of artificial data sets of uncorrelated variables, with two to 10 predictor variables and one dependent variable. We then compared results from stepwise regression with a regression model in which all predictor variables were entered simultaneously. These analyses clearly demonstrate that significance tests based on stepwise procedures lead to greatly inflated Type I error rates (i.e., the probability of erroneously rejecting a true null hypothesis). By using a simple simulation design, our study amplifies previous warnings about using stepwise procedures, and we follow others in recommending that biologists refrain from applying these methods.

Discriminant function analysis with nonindependent data : consequences and an alternative

The discriminant function analysis (DFA) is a multivariate method that is frequently used in bioacoustic research to examine, for instance, whether calls from different species, contexts, or social groups can be distinguished by their acoustic properties. Most published studies include more than one call per subject into such an analysis. This, in fact, leads to a two-factorial data set that includes the factor ‘subject’ in addition to the factor of interest (e.g. species, context, or social group). The regular version of the DFA, however, does not allow for the analysis of such data sets without violating the assumption of independence. In this paper, we show that analysing factorial data sets using a conventional DFA is a case of pseudoreplication and tends to produce (sometimes grossly) incorrect results. In such a case the discriminability of species, contexts or groups etc. can be drastically overestimated. Furthermore, we provide a permutation-based procedure that copes with such data sets.

Long-term reciprocation of grooming in wild West African chimpanzees

Humans are well known for their ability to keep track of social debts over extended periods of time, and for their tendency to preferentially cooperate with closely bonded partners. Non-human primates have been shown to cooperate with kin and non-kin, and reciprocate helpful acts. However, there is ongoing debate over whether they keep track of previous interactions and, if so, whether they can do it over extended periods of time, or are constrained to finalize exchanges within a single encounter. In this study, we used 3000 hours of all-day focal follows of wild chimpanzees (Pan troglodytes verus) to investigate whether both females and males reciprocate grooming within a single interaction, throughout the day, or over longer periods of time. We found that grooming was reciprocated more symmetrically when measured on a long-term, rather than on an immediate or short-term basis. Random giving, general allocation of grooming efforts, similarities among individuals and kinship do not appear to explain these highly reciprocal exchanges. Previously collected consecutive focal follows of single individuals revealed that dyads groomed an average of once every 7 days. Our findings strongly suggest that chimpanzees, similar to humans, are able to keep track of past social interactions, at least for a one-week period, and balance services over repeated encounters.

Lethal aggression in Pan is better explained by adaptive strategies than human impacts

Observations of chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) provide valuable comparative data for understanding the significance of conspecific killing. Two kinds of hypothesis have been proposed. Lethal violence is sometimes concluded to be the result of adaptive strategies, such that killers ultimately gain fitness benefits by increasing their access to resources such as food or mates. Alternatively, it could be a non-adaptive result of human impacts, such as habitat change or food provisioning. To discriminate between these hypotheses we compiled information from 18 chimpanzee communities and 4 bonobo communities studied over five decades. Our data include 152 killings (n = 58 observed, 41 inferred, and 53 suspected killings) by chimpanzees in 15 communities and one suspected killing by bonobos. We found that males were the most frequent attackers (92% of participants) and victims (73%); most killings (66%) involved intercommunity attacks; and attackers greatly outnumbered their victims (median 8:1 ratio). Variation in killing rates was unrelated to measures of human impacts. Our results are compatible with previously proposed adaptive explanations for killing by chimpanzees, whereas the human impact hypothesis is not supported.

Mothers matter! Maternal support, dominance status and mating success in male bonobos (Pan paniscus)

Variation in male mating success is often related to rank differences. Males who are unable to monopolize oestrous females alone may engage in coalitions, thus enhancing their mating success. While studies on chimpanzees and dolphins suggest that coalitions are independent of kinship, information from female philopatric species shows the importance of kin support, especially from mothers, on the reproductive success of females. Therefore, one might expect a similar effect on sons in male philopatric species. We evaluate mating success determinants in male bonobos using data from nine male individuals from a wild population. Results reveal a steep, linear male dominance hierarchy and a positive correlation between dominance status and mating success. In addition to rank, the presence of mothers enhances the mating success of sons and reduces the proportion of matings by the highest ranking male. Mothers and sons have high association rates and mothers provide agonistic aid to sons in conflicts with other males. As bonobos are male-philopatric and adult females occupy high dominance status, maternal support extends into adulthood and females have the leverage to intervene in male conflicts. The absence of female support to unrelated males suggests that mothers gain indirect fitness benefits by supporting their sons.

Diving seabirds share foraging space and time within and among species

Ecological theory predicts that animals with similar foraging strategies should not be able to co-exist without segregating either in space, time or diet. In communities, intra-specific competition is thought to be more intense than the competition among species, because of the lack of niche partitioning between conspecifics. Hence, while different seabird species can overlap in their foraging distribution, intra-specific competition can drive the neighboring populations of the same species to spatial segregation of foraging areas. To investigate ecological segregation within and among species of diving seabirds, we used a multi-species GPS-tracking approach of seabirds of four species on a small island in the Southwest Atlantic. The present study goes beyond previous work by analyzing simultaneous effects of species and colonies. We observed strikingly strong spatial foraging segregation among birds of the same species, breeding in colonies as close as 2 km from each other. Conspecifics from neighboring colonies used foraging places adjacent to their own colony, and there was little or no overlap with birds from the other colony. A zone with increased predator concentration was completely avoided during foraging trips, likely contributing to the spatial segregation. In addition to spatial segregation, we also observed intra-specific differences in other components of foraging behavior, such as time of day, dive depth and diet. These were most likely caused by optimal foraging of individuals in relation to habitat differences on a local scale, leading to a complex pattern of interactions with environmental covariates, in particular foraging daytime, foraging water layer temperature and depth, distance to coast and bathymetric depth of foraging areas. As mechanisms leading to the spatial segregation we propose a combination of optimal foraging and avoidance of predation.

Song repertoire size is correlated with body measures and arrival date in common nightingales, Luscinia megarhynchos

In songbirds, the individual features of a males song play a role in female choice and male–male interactions. Therefore, it has been hypothesized that the song characteristics of an individual should be correlated with its quality and this has been shown for some species. The species studied so far have only small or medium-sized song repertoires, leaving open the question of whether, in species with large repertoires, song characteristics such as repertoire size are related to male quality. We investigated the correlations between body measures, song characteristics and arrival dates of common nightingales, a songbird species in which males have extraordinarily large song type repertoires. We found that birds with large repertoires had longer wings and were heavier, and arrived earlier, than those with small repertoires. Other song characteristics, such as song diversity and proportion of whistle songs, were not significantly correlated with body measures. When we correlated body measures with arrival date, we found that birds that arrived earlier were heavier and tended to be in better condition. These findings provide supporting evidence that repertoire size is an indicator of a males quality.

Issues in information theory-based statistical inference—a commentary from a frequentist’s perspective

After several decades during which applied statistical inference in research on animal behaviour and behavioural ecology has been heavily dominated by null hypothesis significance testing (NHST), a new approach based on information theoretic (IT) criteria has recently become increasingly popular, and occasionally, it has been considered to be generally superior to conventional NHST. In this commentary, I discuss some limitations the IT-based method may have under certain circumstances. In addition, I reviewed some recent articles published in the fields of animal behaviour and behavioural ecology and point to some common failures, misunderstandings and issues frequently appearing in the practical application of IT-based methods. Based on this, I give some hints about how to avoid common pitfalls in the application of IT-based inference, when to choose one or the other approach and discuss under which circumstances a mixing of the two approaches might be appropriate.

Geographical and seasonal variation of harbour porpoise ( Phocoena phocoena ) presence in the German Baltic Sea revealed by passive acoustic monitoring

The harbour porpoise is the only resident cetacean species in the German Baltic Sea. Within the last several decades this harbour porpoise stock declined drastically, causing deep concern about its status. Plans of the German government for proposing Marine Protected Areas (MPAs) to implement Natura 2000 and for assessing the impact of offshore windmill constructions on the marine environment led to an increased research effort on the harbour porpoise in German waters. For the first time, long-term passive acoustic monitoring has been conducted in the German Baltic Sea from the Kiel Bight to the Pomeranian Bay from August 2002 to December 2005. Porpoise detectors (T-PODs) have been installed five to seven metres below the water surface at up to 42 measuring positions throughout the investigated area, registering the exact times of echolocation signals of passing harbour porpoises. The proportion of monitored days with porpoise detection in each quarter of the years has been analysed. A correlation of the results with the longitude of the measuring position revealed a significant decrease from west to east in the percentage of days with porpoise detections. Comparison of data gathered in the first quarters with the third quarters of the monitoring years displayed a seasonal variation with fewer days of porpoise detections in winter time than in summer time. Nevertheless, harbour porpoises have been detected year-round at most of the measuring positions in the German Baltic Sea. The present study clearly indicates a regular use of the German Baltic Sea by harbour porpoises with a geographical and seasonal variation in the usage of the German Baltic Sea. The larger numbers of harbour porpoise detections in spring to autumn compared with winter suggests that the German Baltic Sea is an important breeding and mating area for these animals.

Contingent cooperation between wild female baboons

The apparent rarity of contingent cooperation in animals has convinced many investigators that such reciprocity is unimportant, stimulating consideration of alternative explanations for cooperation, such as by-product mutualism and biological markets motivated by the likelihood of immediate reward. Nevertheless, there is also limited evidence that animals do sometimes rely on memory of recent interactions when behaving altruistically toward others. Here we describe a playback experiment conducted on wild female baboons, suggesting that contingent cooperation may occur among unrelated individuals, even when there is a temporal delay between the two cooperative acts. Hearing the recruitment call of an unrelated recent grooming partner caused subjects to move in the direction of the loudspeaker and approach their former partner, particularly when this partner had an infant. When the subject and her partner were close kin no such effect was observed. Subjects’ responses were not influenced by any type of recent interaction, because prior grooming and prior aggression influenced their behavior in qualitatively different ways. Similarly, their responses were not prompted only by the motivation to resume friendly interactions, because prior grooming alone did not elicit approach. Instead, subjects were most likely to approach their former grooming partner when they had also heard her recruitment call. Results suggest that at least some forms of cooperation in animals may be based on memory of specific recent interactions.