Changes in interspecies association patterns of Atlantic bottlenose dolphins,Tursiops truncatus,and Atlantic spotted dolphins,Stenella frontalis,after demographic changes related to environmental disturbance

Animal populations can be affected by environmental disturbances in many ways including demographic and behavioral changes. This can affect interspecies associations for regularly interacting sympatric species, like bottlenose and spotted dolphins in the Bahamas (observed since 1985 and interspecies associations analyzed since 1993). After two hurricanes in 2004 each species lost roughly 30% of their respective communities resulting in differing social structure and behavioral changes. During mixed species encounters (MSE) group sizes for spotted dolphins ( = 14.1 ± 9.2) were significantly larger than bottlenose dolphins ( = 6.0 ± 7.3; F = 11.74, df = 1, P < 0.001), however, t‐tests revealed no differences between aggressive vs. affiliative encounters. Sexual/aggressive behavior regularly seen previously was not observed posthurricanes and aggressive encounters were greatly reduced. Generally results were similar to prehurricane data including high resightings of spotted dolphins with male alliances prevalent (including new juvenile alliances seen only posthurricane), and individualized bottlenose participation with few male alliances. However temporal associations varied compared to prehurricane. Interspecies association and behavior patterns were altered and likely affected by the changes in intraspecies association patterns following the hurricanes. However both species still participated in MSE, suggesting this is an important component of their ability to coexist as sympatric species.     

Community structure and cluster definition of Atlantic spotted dolphins,Stenella frontalis,in the Bahamas

Fission‐fusion dynamics typical of many delphinid populations allow for a variety of social grouping patterns. Identifying these groupings is crucial before conducting a detailed social structure analysis. This study analyzed the structure of a population of Bahamian spotted dolphins, Stenella frontalis. Through long‐term observations and preliminary analysis, three clusters were defined: Northern, Central, and Southern. To quantitatively investigate these delineations, we conducted analysis on 12 yr of sighting data using SOCPROG 2.3. Coefficients of association (CoA) were calculated using the half‐weight index, with individuals sighted six or more times per pooled period (3 yr each). Nonmetric multidimensional scaling (MD), hierarchical agglomerative cluster analysis and Mantel tests were conducted to determine if any divisions were present. Mantel tests and MD plots analysis supported the delineations into the three clusters. Cluster analysis showed cluster groupings, but with less clear distinctions between the clusters. The amount and strength of associations were significantly higher within clusters than between clusters. Based on behavioral and geographic overlap, these clusters did not meet the definition of separate communities and thus were termed social clusters. These fine scale, within community divisions, are biologically and socially important aspects of their community and are crucial in understanding the dolphins’ social structure.     

Long‐term social structure of a resident community of Atlantic spotted dolphins,Stenella Frontalis,in the Bahamas 1991–2002

Long‐term social structure data on small delphinids is lacking for most species except the bottlenose dolphin. This study describes the long‐term social structure of one community of Atlantic spotted dolphins, Stenella frontalis, divided into three social clusters. Data from 12 yr were analyzed using SOCPROG 2.3. Coefficients of association (CoA) were calculated using the half‐weight index. The overall mean community CoA ranged from 0.09 to 0.12. Temporal analyses and mantel tests revealed significant differences between sex class associations due to high male‐male CoA (0.12–0.23) compared to female‐female and mixed sex CoA (0.08–0.10). Female associations were strongly influenced by reproductive status, calf care, and social familiarity, but not by age class. Male associations were strongly influenced by age, access to females, and alliance formation. Males showed two levels of alliance formation, long‐term first order pairs/trios (CoA 0.70–1.00) and shorter‐term second order alliances between two or more first order alliances (CoA 0.45–0.69), and a possible third level during interspecies interactions. Mating strategies, sex, and cluster formation shaped the social structure in this spotted dolphin community. Similar to many bottlenose dolphin studies, long‐term affiliations for spotted dolphins were correlated with age, sex, and reproductive status.     

Biphonal calls in Atlantic spotted dolphins (Stenella frontalis): bitonal and burst-pulse whistles

Biphonation, the simultaneous production of two sounds by a single animal, is found in the vocalizations of a range of mammalian species. Its prevalence suggests it plays an important role in acoustic communication. Concurrent vocal and behavioural recordings were made of Atlantic spotted dolphins (Stenella frontalis) off Bimini, The Bahamas. The occurrence of two types of biphonal signals is reported: burst-pulse whistles with combined tonal and burst-pulse elements, and bitonal whistles. Biphonal whistles are rarely described in reports of dolphin acoustic repertoires, but were common in these dolphins: of all whistles analysed (n = 1211), 26.84% were burst-pulse whistles and 4.71% were bitonal whistles. A subset of whistles (n = 397) were attributed to dolphins of specific age classes, and used to compare prevalence of biphonation across age. Biphonation occurred in 61.54% of sexually mature and 48.32% of sexually immature dolphins’ whistles. Sexually immature dolphins emitted more burst-pulse whistles than older dolphins: 44.13% of sexually immature dolphins’ whistles were burst-pulse whistles, while 15.38% of adult whistles were burst-pulse whistles. Bitonal whistle production was more prevalent in sexually mature dolphins: 41.03% of adult whistles were bitonal, while only 4.19% of sexually immature dolphins’ whistles were bitonal. The prevalence of a biphonal component in specific repeated, stereotyped whistle contours suggests that these acoustic features could be important components of contact calls, or signature whistles. The biphonal components of spotted dolphin whistles may serve to convey additional information as to identity, age or other factors to conspecifics.     

Social communities and spatiotemporal dynamics of association patterns in estuarine bottlenose dolphins

Network analysis has recently been used to delve into the dynamics of cetacean sociality. Few studies, however, have addressed how habitat shape influences sociality, specifically how linear water bodies constrain the space where individuals might interact. We utilized network and spatiotemporal analyses to investigate association patterns and community structure in a bottlenose dolphin population in a linear estuarine system, the Indian River Lagoon (IRL), Florida. Using sighting histories from a multiyear photo‐identification study we examined association patterns for 185 individuals collected over a 6.5 yr period (2002–2008). The population was highly differentiated (S = 0.723) and organized into six distinct social communities (Q = 0.544), spread in an overlapping pattern along the linear system. Social organization differed between communities, with some displaying highly interconnected networks and others comprising loosely affiliated individuals with more ephemeral associations. Temporal patterns indicated short‐term associations were a significant feature of the fission‐fusion dynamics of this population. Spatial analyses revealed that social structure was shaped by an individual's ranging patterns and by social processes including preference and avoidance behavior. Finally, we found that habitat “narrowness” may be a major driving force behind the sociality observed.     

Hematological,serum, and plasma chemical constituents in pantropical spotted dolphins (Stenella attenuata) following chase,encirclement, and tagging

Hematology, serum chemistry, and plasma hormones were evaluated in 72 pantropical spotted dolphins (Stenella attenuata attenuata) from the eastern tropical Pacific in an attempt to define the degree of stress associated with chase and encirclement by a tuna purse seiner, and are here reported for the first time for this species. Dolphins had high levels of dopamine and moderately elevated levels of enzymes indicative of the expected muscle damage following exertion of the chase. The length of time between the start of the capture operation and blood sampling correlated with increases in platelet and white blood cell counts and mean cell hemoglobin concentration, while the length of time between net tie‐down and blood sampling influenced platelet, white blood cell, and eosinophil counts. Ten dolphins recaptured 1–3 d after their first capture had significantly lower serum creatinine kinase, thyroid (T4) and globulin levels compared to values in dolphins sampled at nominal first capture. Although small sample sizes and large individual variation limit interpretation, these data indicate a stress response occurred in all dolphins, but the extent of the response is within the expected range for adaptive responses previously measured in limited numbers of wild mammals.     

Blood mercury concentrations in common bottlenose dolphins from the Indian River Lagoon,Florida: Patterns of social distribution

Social network analysis has been shown to be effective in studying the social structure of cetacean populations. Common bottlenose dolphins (Tursiops truncatus) inhabiting the Indian River Lagoon (IRL), Florida, have among the highest concentrations of total mercury (THg) in blood reported worldwide. The purpose of this study was to examine the relationship between THg concentrations in IRL dolphins and their social affiliations. Whole blood samples from 98 dolphins with photo‐identification sighting histories were collected between 2003–2007 and 2010–2012. Dolphins were categorized into approximate tertiles of low (mean 199.7 μg/L), medium (mean 366.8 μg/L), and high (mean 990.5 μg/L) THg exposure. Social associations between individual dolphins were defined by the proportion of sightings documented with another known individual. Social network measures of individuals and associations between dyads were examined to determine differences among THg categories. Strong social affiliations of individuals within the highest category of THg were found (P = 0.04), suggesting shared exposures among dolphins foraging in specific areas of the estuary. Network measures of strength and affinity were significantly higher in the highest exposure category. This report used social network analysis as a novel way to examine patterns of exposure to an environmental contaminant in a cetacean population.     

Social dynamics and sexual segregation of Australian humpback dolphins (Sousa sahulensis) in Moreton Bay,Queensland

Understanding the social organization of coastal dolphins, particularly those that reside adjacent to major cities and urbanized regions, is important for understanding the vulnerability of populations to human impacts. We investigated the sexual segregation and social dynamics in a population of Australian humpback dolphins (Sousa sahulensis) inhabiting the near-urban embayment of Moreton Bay, Queensland. Over 160 vessel-based surveys, 148 adults were photographically identified. This included 116 assumed females and 32 assumed males. The population was sexually segregated (SSAS = 0.52, p < .001) and individuals preferred to associate with others of the same sex occupying the same spatial area (p < .025). Association indices between paired males were almost double those of females ($\text{◂=▸}\overline{x}\mathrm{HWI}=0.09$, SD = 0.04 and $\text{◂=▸}\overline{x}\mathrm{HWI}=0.05$, SD = 0.02, respectively). However, females were more gregarious than males and maintained a larger network of associates based on preferred companionships. The social network was also modular with five clusters identified, and a limited number of individuals had a disproportionate influence on connectivity between clusters. This highly fragmented social network implies that the population may be vulnerable to social disruptions from human impacts.     

Abundance and site fidelity of Indo‐Pacific Humpback dolphins in the Great Sandy Strait,Queensland, Australia

This study examined the population of Indo‐Pacific humpback dolphins, Sousa chinensis, inhabiting the Great Sandy Strait Marine Park, Queensland, Australia. A total of 106 dolphins were identified during 228 boat‐based surveys, completed between April 2004 and April 2007. Based on the distribution of resighted individuals and the pattern of associations, it was established that this population consists of two largely geographically distinct communities, referred to as the Northern Community (NC) and the Southern Community (SC). The only recorded interaction between the two groups was a single pod composed of one member of the NC and 11 dolphins from the SC. Abundance was estimated for the entire population and by geographical area using open population models. Estimates for the Great Sandy Strait indicate that about 150 dolphins (N_GSS= 148.4, SE = 8.3, 95% CI: 132.5–165.2) used this area during the study. The NC and SC total population sizes was estimated to be 76 (N_NGSS= 75.80, SE = 3.88, 95% CI = 71–86) and 75 (N_SGSS= 74.98, SE = 4.43, 95% CI: 66–83), respectively. Analysis of residence patterns indicates that a majority of the identified dolphins are long‐term residents.     

Common bottlenose dolphin (Tursiops truncatus) social structure and distribution changes following the 2008 Unusual Mortality Event in the Indian River Lagoon,Florida

In animal societies with fission-fusion dynamics, demographic disturbances can influence the social and spatial structure of the population. Within the Indian River Lagoon (IRL), Florida, common bottlenose dolphins (Tursiops truncatus) have experienced recurrent unusual mortality events (UMEs) providing an opportunity to examine postdisturbance population and social cluster restructuring. This study investigates the impact of the potentially nonepizootic 2008 UME on the IRL dolphin population. Photo-identification surveys conducted from August 2006 to May 2010 were stratified into pre- (August 2006–April 2008) and post-UME (September 2008–May 2010) time periods. Social network and spatial (univariate kernel density) analyses were limited to individuals sighted 5+ times per period (pre-UME = 183, post-UME = 134), and indicated a change from 11 to ten social clusters, although individuals did not always reassociate with pre-UME cluster associates. Despite the social and spatial disconnect between IRL proper and Mosquito Lagoon clusters, both network density and core area spatial overlap increased post-UME allowing for increased intercluster interactions. However, intracluster associations increased as well, allowing the population to maintain multiple social clusters within a loosely connected network. This study shows the important role sociality may play in the adaptability of cetaceans to environmental and demographic changes.     

Genetic and family structure in a group of 165 common bottlenose dolphins caught off the Japanese coast

The biological and genetic structure of common bottlenose dolphins (Tursiops truncatus) that migrate seasonally near Japan remains largely unknown. We investigated the genetic and family structure in a group of 165 common bottlenose dolphins caught off the coast of Japan using mitochondrial DNA (mtDNA) and 20 microsatellite DNA markers. Phylogenetic analysis of the mtDNA control region sequences suggested that the dolphins were related more closely to oceanic types from Chinese waters than other geographic regions. The information on sex, sexual maturation and age together with the genetic markers revealed a strong likelihood for 37 familial relationships related mostly to maternity and an under‐representation of juvenile female offspring. The maternal dolphins had a similar offspring‐birth interval as the coastal types from North Atlantic Ocean, but a slightly younger first‐progeny age. The sex bias in the captured group was particularly marked towards an over‐representation of males among the young and immature dolphins, whereas the mature adults had an equal number of males and females. These results should be useful for future comparative biological, genetic and evolutionary investigations of bottlenose dolphins from the North Pacific Ocean with those from other regions.     

Dusky dolphin (Lagenorhynchus obscurus) social structure characterized by social fluidity and preferred companions

Many coastal delphinids exhibit social structures with a high degree of fission‐fusion dynamics. However, little is known about fission‐fusion dynamics in semipelagic delphinids such as dusky dolphins (Lagenorhynchus obscurus). We analyzed dusky dolphin social structure in a winter foraging habitat in New Zealand. We examined the population‐level societal attributes of preferred/avoided companions, behaviorally specific preferred/avoided companions, and simple ratio (SR) association index strength and distribution and the dyad‐level societal attribute of temporal patterning of associations. We identified 741 individuals; 99 met inclusion criteria for analysis. Long‐term and short‐term preferred/avoided companions were present in the pooled (6 yr) and annual samples. Individuals were observed to form long‐term but not short‐term preferred/avoided companions during all behavioral states. Mean and maximum SRs were 0.03 ± 0.01 SD and 0.40 ± 0.20 SD, respectively. Temporal analysis indicated that individuals associated preferentially throughout the study period. Dusky dolphins exhibited high social fluidity, as indicated by many weak and few strong associations. However, there was stability as individuals were observed to form behaviorally specific preferred/avoided companions and associate preferentially throughout the 6 yr period. Dusky dolphins thus display a dynamic social structure where overall low levels of bonding are juxtaposed against a subset of the population that forms strong bonds.     

Status,population trend and genetic structure of South American fur seals,Arctocephalus australis,in southwestern Atlantic waters

The South American fur seal (SAFS), Arctocephalus australis, was severely depleted during the 18th century. This work updates information on distribution, abundance, seasonal movements, and genetic structure along the southwestern South Atlantic. Its distribution in Argentine waters includes nineteen rookeries on the Patagonian coast, Isla de los Estados, and islets of the Beagle Channel. Censuses were carried out in rookeries from Isla Escondida to Isla de los Estados from 1996 to 2014. The largest concentrations are found in Chubut, (4,500–15,500 seals) and Isla de los Estados (4,500). Pups were recorded on Isla Escondida, Rasa, and Isla de los Estados. An increasing number of SAFS were recorded in northern Patagonia, including seals marked in Uruguay. Skin samples were taken from southern Brazil to Tierra del Fuego in order to evaluate population structure and demographic trends. Genetic results indicated no population subdivisions (Φ_st = ?0.0292, F_st = ?0.00716, P > 0.05). Bayesian skyline plots constructed for the entire data set indicated evidence of rapid population expansion discernible in Nef between the last 20,000 and 17,000 yr. Genetics results and observation data from marked fur seals support the hypothesis of ancient gene flow and a single Atlantic population. Therefore, SAFS should be managed under international and integrated conservation policies.     

Seasonal changes in fish catch and environmental variables in a large Tropical Lake,Volta, Ghana

Fish species assemblage and selected environmental variables were monitored in Lake Volta from September 2014 to August 2016 to determine seasonal variability in species composition, catch and environmental variables that determine the structure of the fish community. A total of 1,557 individual fish belonging to 41 species, 25 genera and 13 families were recorded. The important fish species with respect to frequency of occurrence, abundance and weight, respectively, were as follows: Chrysichthys (100%; 43.03%; 17.93%), tilapias (100%; 28.97%; 17.86%), Alestes (100%; 14.13%; 32.10%) and Bagrus (91.67%; 5.65%; 12.80%) in that order. The composition of fish species and their temporal variation in experimental catches were similar to that of commercial catches. There were no significant differences (p > 0.05) in species abundance, weight and diversity indices between the dry and wet seasons. The modal class of length frequency distribution of the dominant species in the catch, Chrysichthys, reduced from 125 mm in 2006 to 95 mm in the current study indicating overfishing. Environmental variables considered showed little variation and within optimal ranges for fish survival and did not differ significantly between seasons. Canonical Correspondence Analysis showed that environmental variables explained 43.30% of the variation in species abundance with Lake water level, nitrite‐nitrogen and total dissolved solids being the main environmental factors influencing the structure of the fish community.     

Bottlenose dolphin (Tursiops truncatus) social structure in the Shannon Estuary,Ireland, is distinguished by age‐ and area‐related associations

Social and network analyses that incorporate information on individuals within a population enhance our knowledge of complex species. In this study, the social structure of wild bottlenose dolphins in the Shannon Estuary, Ireland, was analyzed by examining the dynamics of the whole population and then of individuals classed by sex, age, and area. One hundred and twenty‐one dolphins were identified during 522 sightings between 2012 and 2015. The mean half‐weight association index (HWI) of the population was 0.07 ± 0.02. The highest HWIs for individuals of known sex were for female‐male pairs. Mean within‐class HWI was significantly higher than mean between‐class HWI for both age and area classes. Ordinations and sociograms were used to visualize social networks. Permutation tests revealed nonrandom associations for the population overall and both between and within classes. Temporal analyses showed associations persisting for >1,000 d. The whole population's best fit model was for two levels of casual acquaintances. Movement analyses demonstrated the use of the inner estuary by only 25% of the population revealing a potential community division by area. The difference between mean HWI when socializing (0.09 ± 0.03) compared to foraging (0.06 ± 0.03) was significant. These results highlight the importance of localized research, reflecting the complexity found in bottlenose dolphin societies globally.     

Population structure of Atlantic mackerel inferred from RAD‐seq‐derived SNP markers: effects of sequence clustering parameters and hierarchical SNP selection

Restriction‐site‐associated DNA sequencing (RAD‐seq) and related methods are revolutionizing the field of population genomics in nonmodel organisms as they allow generating an unprecedented number of single nucleotide polymorphisms (SNPs) even when no genomic information is available. Yet, RAD‐seq data analyses rely on assumptions on nature and number of nucleotide variants present in a single locus, the choice of which may lead to an under‐ or overestimated number of SNPs and/or to incorrectly called genotypes. Using the Atlantic mackerel (Scomber scombrus L.) and a close relative, the Atlantic chub mackerel (Scomber colias), as case study, here we explore the sensitivity of population structure inferences to two crucial aspects in RAD‐seq data analysis: the maximum number of mismatches allowed to merge reads into a locus and the relatedness of the individuals used for genotype calling and SNP selection. Our study resolves the population structure of the Atlantic mackerel, but, most importantly, provides insights into the effects of alternative RAD‐seq data analysis strategies on population structure inferences that are directly applicable to other species.     

Factors driving the variability in diving and movement behavior of migrating humpback whales (Megaptera novaeangliae): Implications for anthropogenic disturbance studies

Humpback whales (Megaptera novaeangliae) undertake one of the longest migrations of any animal and while on a broad‐scale this journey appears direct, on a fine‐scale, behaviors associated with socializing and breeding are regularly observed. However, little is known about which social and environmental factors influence behavior during this time. Here we examined the effect of multiple factors on the movement (speed and course) and diving behavior (dive and surfacing duration) of humpback whales during migration off the eastern coast of Australia. Focal data (202 h) were collected on 94 different whale groups with simultaneous social and environmental context data. The environmental factors water depth and wind speed were found to be important predictors of dive and movement behavior, whereas social factors were less influential at this site. Groups tended to dive for longer with increased water depth but traveled more slowly in increasing wind speeds. These baseline studies are crucial when examining the effect of anthropogenic disturbance. Determining which natural factors significantly affect behavior ensures any observed behavioral changes are correctly attributed to the disturbance and are not a result of other factors. In addition, any responses observed can be put into biological context and their relative magnitude determined.