Evidence of male alliance formation in a small dolphin community

The photo-identification of uniquely marked individuals has revealed much about mammalian behaviour and social structure in recent decades. In bottlenose dolphins (Tursiops spp.), for example, the long-term tracking of individuals has unveiled considerable variation in social structure among populations and various spatio-temporal aspects of group formation. In this study, we investigated associations among individual males in a small community of Indo-Pacific bottlenose dolphins (T. aduncus) residing in an urbanized estuary in southwestern Australia. Given the relative proximity of our study area to other populations in which complex male alliances form for the purpose of mate acquisition, we used long-term photo-identification records and social analyses to assess whether such alliances also occur in smaller and more isolated settings. Our work revealed strong social bonds and long-term, non-random associations among individual males, suggesting the occurrence of male alliances. Behavioural observations of alliances interacting with potentially receptive adult females from the estuary community and from adjacent communities, and exhibiting sexual display behaviours near females, suggest that these alliances occur in a reproductive context. As the first formal analysis indicating the occurrence of male alliances outside Shark Bay along the vast western coastline of Australia, this study complements previous research and extends our understanding of the evolutionary and ecological processes that drive alliance formation

Genes or Culture: Are Mitochondrial Genes Associated with Tool Use in Bottlenose Dolphins ( Tursiops sp.)?

Some bottlenose dolphins use marine sponges as foraging tools (‘sponging'), which appears to be socially transmitted from mothers mainly to their female offspring. Yet, explanations alternative to social transmission have been proposed. Firstly, the propensity to engage in sponging might be due to differences in diving ability caused by variation of mitochondrial genes coding for proteins of the respiratory chain. Secondly, the cultural technique of sponging may have selected for changes in these same genes (or other autosomal ones) among its possessors. We tested whether sponging can be predicted by mitochondrial coding genes and whether these genes are under selection. In 29 spongers and 54 non-spongers from two study sites, the non-coding haplotype at the HVRI locus was a significant predictor of sponging, whereas the coding mitochondrial genes were not. There was no evidence of selection in the investigated genes. Our study shows that mitochondrial gene variation is unlikely to be a viable alternative to cultural transmission as a primary driver of tool use in dolphin

Reconstructing the colonization history of Indo-Pacific bottlenose dolphins (Tursiops aduncus) in Northwestern Australia

Bottlenose dolphins (Tursiops spp.) are found in waters around Australia, with T. truncatus typically occupying deeper, more oceanic habitat, while T. aduncus occur in shallower, coastal waters. Little is known about the colonization history of T. aduncus along the Western Australian coastline; however, it has been hypothesized that extant populations are the result of an expansion along the coastline originating from a source in the north of Australia. To investigate the history of coastal T. aduncus populations in the area, we generated a genomic SNP dataset using a double-digest restriction-site-associated DNA (ddRAD) sequencing approach. The resulting dataset consisted of 103,201 biallelic SNPs for 112 individuals which were sampled from eleven coastal and two offshore sites between Shark Bay and Cygnet Bay, Western Australia. Our population genomic analyses showed a pattern consistent with the proposed source in the north with significant isolation by distance along the coastline, as well as a reduction in genomic diversity measures along the coastline with Shark Bay showing the most pronounced reduction. Our demographic analysis indicated that the expansion of T. aduncus along the coastline began around the last glacial maximum and progressed southwards with the Shark Bay population being founded only 13 kya. Our results are in line with coastal colonization histories inferred for Tursiops globally, highlighting the ability of delphinids to rapidly colonize novel coastal niches as habitat is released during glacial cycle-related global sea level and temperature changes

Payoff- and Sex-Biased Social Learning Interact in a Wild Primate Population.

Social learning in animals is now well documented, but few studies have determined the contexts shaping when social learning is deployed. Theoretical studies predict copying of conspecifics gaining higher payoffs [1-4], a bias demonstrated in primates only in captivity [5]. In the wild, research has shown selective attention toward the philopatric sex, a group's stable core [6]. Here, we report the first rigorous experimental test of the existence of a payoff bias in wild primates and its interaction with the sex of the model. We created a payoff bias in which an immigrant alpha male in each of three groups of wild vervet monkeys received five times more food upon opening a foraging box than did the philopatric alpha female, whereas in two control groups, male and female models received the same amount of food. We tested whether this payoff asymmetry would override the previously documented selective learning from resident females. Group members were tested after having watched both models. When both models received the same amount of food, audience members copied the female model significantly more than the male model, confirming previous findings. However, when a marked payoff bias was introduced, male, but not female, vervet monkeys significantly more often copied the male model receiving a higher payoff. These results demonstrate behavioral flexibility in the dispersing sex in these primates and suggest that the philopatric sex can afford to be more conservative in their social learning. Our findings show that multiple social-learning biases can coexist and interact within the same species

Sex bias in biopsy samples collected from free-ranging dolphins

Author Posting. © The Author(s), 2009. This is the author's version of the work. It is posted here by permission of Springer for personal use, not for redistribution. The definitive version was published in European Journal of Wildlife Research 56 (2010): 151-158, doi:10.1007/s10344-009-0299-7.Biological samples of free-ranging dolphins are increasingly used to gain information on population structure and ecology. In small cetaceans, the gender of individuals usually cannot be determined at sea, and population sex ratio has to be inferred indirectly. We used molecular sexing to determine the gender of 340 biopsy samples of bottlenose dolphins, Tursiops truncatus, spotted dolphins, Stenella frontalis, and common dolphins, Delphinus delphis, collected around the Azores and Madeira. Sex ratio was globally skewed in favor of males, and differed between species and archipelagos. Skew was probably influenced by the selectivity of biopsy collectors and seasonal or year-round predominance of males in natural populations. Skew was also influenced by sampling duration and intensity. In the Azores, when several samples were successively collected within the same group, the proportion of female samples decreased as a function of sample order. This trend indicated a tendency for females to increasingly avoid the boat while samples were being collected. It showed that males and females reacted differently to the perturbation caused by the biopsy sampling process (i.e. sample collection and driving style).Portuguese Foundation for Science and Technology (FCT) and the FEDER program for funding the CETAMARH (POCTI/BSE/38991/01) and the GOLFINICHO (POCI/BIA-BDE/61009/2004) projects, S.Q.'s post-doctoral grants (IMAR/FCT- PDOC-006/2001-MoleGen and SFRH/BPD/19680/2004), M.A.S.'s doctoral (SFRH/BD/8609/2002) and post-doctoral (SFRH/BPD/29841/2006) grants, S.M.'s investigation assistant grant (CETAMARHII/POCTI/BSE/38991/2001) and I.C.'s investigation assistant grants (IMAR/FCT/GOLFINICHO/001/2005 and IMAR/FCT/GOLFINICHO/004/2006). FCT for its pluri-annual funding to Research Unit #531 and the EU funded program Interreg IIIb for funding the MACETUS project (MAC/4.2/M10) as well as R.P. and S.M.’s grants (IMAR/INTERREGIIIb/MACETUS/MAC1/2)

Thar She Blows! A Novel Method for DNA Collection from Cetacean Blow

Background: Molecular tools are now widely used to address crucial management and conservation questions. To date, dart biopsying has been the most commonly used method for collecting genetic data from cetaceans; however, this method has some drawbacks. Dart biopsying is considered inappropriate for young animals and has recently come under scrutiny from ethical boards, conservationists, and the general public. Thus, identifying alternative genetic collection techniques for cetaceans remains a priority, especially for internationally protected species. Methodology/Principal Findings: In this study, we investigated whether blow-sampling, which involves collecting exhalations from the blowholes of cetaceans, could be developed as a new less invasive method for DNA collection. Our current methodology was developed using six bottlenose dolphins, Tursiops truncatus, housed at the National Aquarium, Baltimore (USA), from which we were able to collect both blow and blood samples. For all six individuals, we found that their mitochondrial and microsatellite DNA profile taken from blow, matched their corresponding mitochondrial and microsatellite DNA profile collected from blood. This indicates that blow-sampling is a viable alternative method for DNA collection. Conclusion/Significance: In this study, we show that blow-sampling provides a viable and less invasive method for collection of genetic data, even for small cetaceans. In contrast to dart biopsying, the advantage of this method is that it capitalizes on the natural breathing behaviour of dolphins and can be applied to even very young dolphins. Both biopsy and blow-sampling require close proximity of the boat, but blow-sampling can be achieved when dolphins voluntarily bowride and involves no harmful contact

Tool use and social homophily among male bottlenose dolphins

Homophilous behaviour plays a central role in the formation of human friendships. Individuals form social ties with others that show similar phe-notypic traits, independently of relatedness. Evidence of such homophily can be found in bottlenose dolphins (Tursiops aduncus) in Shark Bay, Western Australia, where females that use marine sponges as foraging tools often associate with other females that use sponges. 'Sponging' is a socially learned, time-consuming behaviour, transmitted from mother to calf. Previous research illustrated a strong female bias in adopting this technique. The lower propensity for males to engage in sponging may be due to its incompatibility with adult male-specific behaviours, particularly the formation of multi-level alliances. However, the link between sponging and male behaviour has never been formally tested. Here, we show that male spongers associated significantly more often with other male spongers irrespective of their level of relatedness. Male spongers spent significantly more time foraging, and less time resting and travelling, than did male non-spongers. Interestingly, we found no difference in time spent socializing. Our study provides novel insights into the relationship between tool use and activity budgets of male dolphins, and indicates social homophily in the second-order alliance composition of tool-using bottlenose dolphins

Why Do Dolphins Carry Sponges?

Tool use is rare in wild animals, but of widespread interest because of its relationship to animal cognition, social learning and culture. Despite such attention, quantifying the costs and benefits of tool use has been difficult, largely because if tool use occurs, all population members typically exhibit the behavior. In Shark Bay, Australia, only a subset of the bottlenose dolphin population uses marine sponges as tools, providing an opportunity to assess both proximate and ultimate costs and benefits and document patterns of transmission. We compared sponge-carrying (sponger) females to non-sponge-carrying (non-sponger) females and show that spongers were more solitary, spent more time in deep water channel habitats, dived for longer durations, and devoted more time to foraging than non-spongers; and, even with these potential proximate costs, calving success of sponger females was not significantly different from non-spongers. We also show a clear female-bias in the ontogeny of sponging. With a solitary lifestyle, specialization, and high foraging demands, spongers used tools more than any non-human animal. We suggest that the ecological, social, and developmental mechanisms involved likely (1) help explain the high intrapopulation variation in female behaviour, (2) indicate tradeoffs (e.g., time allocation) between ecological and social factors and, (3) constrain the spread of this innovation to primarily vertical transmission

The Use of Carcasses for the Analysis of Cetacean Population Genetic Structure: A Comparative Study in Two Dolphin Species

Advances in molecular techniques have enabled the study of genetic diversity and population structure in many different contexts. Studies that assess the genetic structure of cetacean populations often use biopsy samples from free-ranging individuals and tissue samples from stranded animals or individuals that became entangled in fishery or aquaculture equipment. This leads to the question of how representative the location of a stranded or entangled animal is with respect to its natural range, and whether similar results would be obtained when comparing carcass samples with samples from free-ranging individuals in studies of population structure. Here we use tissue samples from carcasses of dolphins that stranded or died as a result of bycatch in South Australia to investigate spatial population structure in two species: coastal bottlenose (Tursiops sp.) and short-beaked common dolphins (Delphinus delphis). We compare these results with those previously obtained from biopsy sampled free-ranging dolphins in the same area to test whether carcass samples yield similar patterns of genetic variability and population structure. Data from dolphin carcasses were gathered using seven microsatellite markers and a fragment of the mitochondrial DNA control region. Analyses based on carcass samples alone failed to detect genetic structure in Tursiops sp., a species previously shown to exhibit restricted dispersal and moderate genetic differentiation across a small spatial scale in this region. However, genetic structure was correctly inferred in D. delphis, a species previously shown to have reduced genetic structure over a similar geographic area. We propose that in the absence of corroborating data, and when population structure is assessed over relatively small spatial scales, the sole use of carcasses may lead to an underestimate of genetic differentiation. This can lead to a failure in identifying management units for conservation. Therefore, this risk should be carefully assessed when planning population genetic studies of cetaceans

A New Dolphin Species, the Burrunan Dolphin Tursiops australis sp. nov., Endemic to Southern Australian Coastal Waters

Small coastal dolphins endemic to south-eastern Australia have variously been assigned to described species Tursiops truncatus, T. aduncus or T. maugeanus; however the specific affinities of these animals is controversial and have recently been questioned. Historically ‘the southern Australian Tursiops’ was identified as unique and was formally named Tursiops maugeanus but was later synonymised with T. truncatus. Morphologically, these coastal dolphins share some characters with both aforementioned recognised Tursiops species, but they also possess unique characters not found in either. Recent mtDNA and microsatellite genetic evidence indicates deep evolutionary divergence between this dolphin and the two currently recognised Tursiops species. However, in accordance with the recommendations of the Workshop on Cetacean Systematics, and the Unified Species Concept the use of molecular evidence alone is inadequate for describing new species. Here we describe the macro-morphological, colouration and cranial characters of these animals, assess the available and new genetic data, and conclude that multiple lines of evidence clearly indicate a new species of dolphin. We demonstrate that the syntype material of T. maugeanus comprises two different species, one of which is the historical ‘southern form of Tursiops’ most similar to T. truncatus, and the other is representative of the new species and requires formal classification. These dolphins are here described as Tursiops australis sp. nov., with the common name of ‘Burrunan Dolphin’ following Australian aboriginal narrative. The recognition of T. australis sp. nov. is particularly significant given the endemism of this new species to a small geographic region of southern and south-eastern Australia, where only two small resident populations in close proximity to a major urban and agricultural centre are known, giving them a high conservation value and making them susceptible to numerous anthropogenic threats