Benefits of living closer to kin vary by genealogical relationship in a territorial mammal.

While cooperative interactions among kin are a key building block in the societies of group-living species, their importance for species with more variable social environments is unclear. North American red squirrels (Tamiasciurus hudsonicus) defend individual territories in dynamic neighbourhoods and are known to benefit from living among familiar conspecifics, but not relatives. However, kin-directed behaviours may be restricted to specific genealogical relationships or strongly mediated by geographical distance, masking their influence at broader scales. Using distance between territories as a proxy for the ability of individuals to interact, we estimated the influence of primary kin (parents, offspring, siblings) on the annual survival and reproductive success of red squirrels. This approach revealed associations between fitness and access to kin, but only for certain genealogical relationships and fitness components. For example, females had enhanced annual survival when living closer to their daughters, though the reverse was not true. Most surprising was the finding that males had higher annual reproductive success when living closer to their father, suggesting possible recognition and cooperation among fathers and sons. Together, these findings point to unexpected nuance in the fitness consequences of kinship dynamics for a species that is territorial and largely solitary.

Causation, not collinearity: Identifying sources of bias when modelling the evolution of brain size and other allometric traits.

Many biological traits covary with body size, resulting in an allometric relationship. Identifying the evolutionary drivers of these traits is complicated by possible relationships between a candidate selective agent and body size itself, motivating the widespread use of multiple regression analysis. However, the possibility that multiple regression may generate misleading estimates when predictor variables are correlated has recently received much attention. Here, we argue that a primary source of such bias is the failure to account for the complex causal structures underlying brains, bodies, and agents. When brains and bodies are expected to evolve in a correlated manner over and above the effects of specific agents of selection, neither simple nor multiple regression will identify the true causal effect of an agent on brain size. This problem results from the inclusion of a predictor variable in a regression analysis that is (in part) a consequence of the response variable. We demonstrate these biases with examples and derive estimators to identify causal relationships when traits evolve as a function of an existing allometry. Model mis-specification relative to plausible causal structures, not collinearity, requires further consideration as an important source of bias in comparative analyses.

Cross-modal perception of identity by sound and taste in bottlenose dolphins.

While studies have demonstrated concept formation in animals, only humans are known to label concepts to use them in mental simulations or predictions. To investigate whether other animals use labels comparably, we studied cross-modal, individual recognition in bottlenose dolphins (Tursiops truncatus) that use signature whistles as labels for conspecifics in their own communication. First, we tested whether dolphins could use gustatory stimuli and found that they could distinguish between water and urine samples, as well as between urine from familiar and unfamiliar individuals. Then, we paired playbacks of signature whistles of known animals with urine samples from either the same dolphin or a different, familiar animal. Dolphins investigated the presentation area longer when the acoustic and gustatory sample matched than when they mismatched. This demonstrates that dolphins recognize other individuals by gustation alone and can integrate information from acoustic and taste inputs indicating a modality independent, labeled concept for known conspecifics.

Species information in whistle frequency modulation patterns of common dolphins.

The most flexible communication systems are those of open-ended vocal learners that can acquire new signals throughout their lifetimes. While acoustic signals carry information in general voice features that affect all of an individual's vocalizations, vocal learners can also introduce novel call types to their repertoires. Delphinids are known for using such learned call types in individual recognition, but their role in other contexts is less clear. We investigated the whistles of two closely related, sympatric common dolphin species, Delphinus delphis and Delphinus bairdii, to evaluate species differences in whistle contours. Acoustic recordings of single-species groups were obtained from the Southern California Bight. We used an unsupervised neural network to categorize whistles and compared the resulting whistle types between species. Of the whistle types recorded in more than one encounter, 169 were shared between species and 60 were species-specific (32 D. delphis types, 28 D. bairdii types). Delphinus delphis used 15 whistle types with an oscillatory frequency contour while only one such type was found in D. bairdii. Given the role of vocal learning in delphinid vocalizations, we argue that these differences in whistle production are probably culturally driven and could help facilitate species recognition between Delphinus species. This article is part of the theme issue 'Vocal learning in animals and humans'.

Vocal sequences in narwhals (Monodon monoceros).

Sequences are indicative of signal complexity in vocal communication. While vocal sequences are well-described in birds and terrestrial mammals, the extent to which marine mammals use them is less well understood. This study documents the first known examples of sequence use in the narwhal (Monodon monoceros), a gregarious Arctic cetacean. Eight female narwhals were fitted with animal-borne recording devices, resulting in one of the largest datasets of narwhal acoustic behaviour to date. A combination of visual and quantitative classification procedures was used to test whether subjectively defined vocalization patterns were organized into sequences. Next, acoustic characteristics were analyzed to assess whether sequences could disclose group or individual identity. Finally, generalized linear models were used to investigate the behavioural context under which sequences were produced. Two types of sequences, consisting of "paired" patterns and "burst pulse series," were identified. Sequences of burst pulse series were typically produced in periods of high vocal activity, whereas the opposite was true for sequences of paired patterns, suggesting different functions for each. These findings extend the set of odontocetes which are known to use vocal sequences. Inquiry into vocal sequences in other understudied marine mammals may provide further insights into the evolution of vocal communication.

Repeated downsweep vocalizations of the Araguaian river dolphin, Inia araguaiaensis.

Araguaian botos (Inia araguaiaensis) are known to produce pulsed as well as tonal sounds. This study documents the first evidence for repetitive sequences of downsweep whistles in botos that appear to be shared between individuals, and the context of their occurrence is investigated. Boat surveys were conducted along the Tocantins River located in the Eastern Amazon over a period of 42 days between 2012 and 2018. Eighty-two groups of Araguaian botos were observed, and 43 h of sound recordings were acquired. 632 downsweep whistles were recorded in 10 encounters. Four of these encounters contained downsweep bouts (21 bouts with ≥2 whistles) with short inter-call intervals (bout criterion 50 s) and up to 161 whistles. A statistical relationship was not found between downsweep occurrence and any of the contextual parameters that were investigated, including socializing, travelling, feeding, group size, presence of calves, and socio-sexual displays. The rarity of these signals makes them unlikely candidates for individual or group identification. It is more likely that they are associated with very specific contexts, such as nursing or mating, both of which were rarely observed in this study. Further studies are required to investigate context specificity and elucidate the function of these signals.