Broad-scale impacts of coastal mega-infrastructure project on obligatory inshore delphinids: A cautionary tale from Hong Kong.

Inshore marine habitats experience considerable anthropogenic pressure, as this is where many adverse effects of human activities concentrate. In the rapidly-changing seascape of the Anthropocene, Hong Kong waters at the heart of world's fastest developing coastal region can serve as a preview-window into coastal seas of the future, with ever-growing anthropogenic footprint. Here, we quantify how large-scale coastal infrastructure projects can affect obligatory inshore cetaceans, bringing about population-level consequences that may compromise their long-term demographic viability. As a case in point, we look at the construction of world's longest sea crossing system and broad-scale demographic, social and spatial responses it has caused in a shallow-water delphinid, the Indo-Pacific humpback dolphin (Sousa chinensis). Soon after the infrastructure project began, dolphins markedly altered their home range near construction sites such that these waters no longer functioned as dolphin core areas despite the apparent presence of prey, indicating that anthropogenic impacts outweighed foraging benefits. The contraction of key habitats has in turn led individuals to interact over spatially more constricted area, reshaping their group dynamics and social network. Although there was no apparent decline in dolphin numbers that could be detected with mark-recapture estimates, adult survival rates decreased drastically from 0.960 to 0.904, the lowest estimate for these animals anywhere across the region to date, notably below the previously estimated demographic threshold of their long-term persistence (0.955). It is apparent that during an advanced stage of this coastal infrastructure project, dolphins were under a major anthropogenic pressure that, if sustained, could be detrimental to their long-term persistence as a viable demographic unit. As effective conservation of species and habitats depends on informed management decisions, this study offers a valuable lesson in environmental risk assessment, underscoring the implications of human-induced rapid environmental change on obligatory inshore delphinids-sentinels of coastal habitats that are increasingly degraded in fast-changing coastal seas.

Prey decline leads to diet shift in the largest population of Indo-Pacific humpback dolphins?

The Pearl River Delta (PRD) region on the southeast coast of China has long been known as a highly productive fishing ground. Since the late 1980s, fishing pressure in the PRD has been intense, which warrants concerns of potential fishery-related impacts on the food resources and foraging ecology of apex marine predators in this region, such as the Indo-Pacific humpback dolphin (Sousa chinensis). In this study, we examined 54 stomachs with food remains, collected from beached carcasses of humpback dolphins recovered during fifteen years between 2003 and 2017. The 6043 identified prey items represent 62 teleost taxa, primarily small estuarine fish, but also larger reef fish. The dolphins appear to be opportunistic foragers, hunting across the water-column, with preference for shoaling and meaty fishes (e.g. Collichthys lucidus IRI% = 38.6%, Johnius belangerii IRI% = 23.1%, Mugil cephalus IRI% = 14.0%). Our findings suggest a dietary shift in recent years, from primarily demersal (as previously reported) to greater intake of neritic and pelagic fish. Dolphin foraging group size has decreased in recent years, which corresponds with declining size and numbers of prey items retrieved from dolphin stomachs. We suggest that these are indicators of declining food resources. Faced with a shortage of preferred prey, humpback dolphins may have broadened their dietary spectrum to maintain their daily energy intake, while their foraging group size decreased in response to the altered tradeoff between the costs and benefits of group foraging.

Photo-identification comparison of four Indo-Pacific humpback dolphin populations off southeast China.

Indo-Pacific humpback dolphins (Sousa chinensis) inhabit shallow coastal waters of the Indo-Pacific region including southeast China, with at least 6 putative populations identified to date in Chinese waters. However, the connectivity among these populations has not yet been fully investigated. In the present study, we compared and cross-matched photographic catalogs of individual dolphins collected to date in the Pearl River Delta region, Leizhou Bay, Sanniang Bay, and waters southwest of Hainan Island, a total of 3158 individuals, and found no re-sighting of individual dolphins among the 4 study areas. Furthermore, there was a notable difference in the pigmentation pattern displayed by individuals from these 4 regions. We suggest that this may be a phenotypical expression of fine-scale regional differentiation among humpback dolphin groups, possibly distinct populations. Given the considerable conservation management implications it may carry (e.g. definition of management units), further research is much needed.

Genomics of Population Differentiation in Humpback Dolphins, Sousa spp. in the Indo-Pacific Ocean.

Speciation is a fundamental process in evolution and crucial to the formation of biodiversity. It is a continuous and complex process, which can involve multiple interacting barriers leading to heterogeneous genomic landscapes with various peaks of divergence among populations. In this study, we used a population genomics approach to gain insights on the speciation process and to understand the population structure within the genus Sousa across its distribution in the Indo-Pacific region. We found 5 distinct clusters, corresponding to S. plumbea along the eastern African coast and the Arabian Sea, the Bangladesh population, S. chinensis off Thailand and S. sahulensis off Australian waters. We suggest that the high level of differentiation found, even across geographically close areas, is likely determined by different oceanographic features such as sea surface temperature and primary productivity.

Epidermal Lesions and Injuries of Coastal Dolphins as Indicators of Ecological Health.

Humpback dolphins (genus Sousa), obligatory inshore delphinids, are frequently exposed to adverse effects of many human activities. In Hong Kong, one of the world's most urbanised coastal regions, ~ 50% of the dolphins suffer from at least one type of epidermal lesions, likely related to anthropogenically degraded habitat. Furthermore, one in every ten dolphins has physical injuries indicative of vessel collisions, propeller cuts and fishing-gear entanglements. As top predators with long lifespan, dolphins are good "barometers" of marine environment and their compromised health conditions are symptomatic of increasingly degraded ecological conditions of coastal seas, especially in rapidly developing regions of fast-growing economies.

Patterns of geographic variation between mitochondrial and nuclear markers in Heaviside's (Benguela) dolphins (Cephalorhynchus heavisidii).

The Heaviside's (or Benguela) dolphin (Cephalorhynchus heavisidii) is endemic to the west coast of southern Africa. The present study investigated the population genetic structure across a large portion of the species distribution using mitochondrial control region and nuclear (microsatellite) markers. A total of 395 biopsy skin samples were analyzed; they were collected from free-ranging Heaviside's dolphins in 7 locations along 1650 km of coast between Table Bay, South Africa and Walvis Bay, Namibia. Both genetic markers rejected the hypothesis of 1 homogenous population but revealed contrasting results in the genetic structuring of putative populations. Mitochondrial DNA suggested either 2 populations or a fine-scale division with 6 (sub) populations, while microsatellite markers were indicative of 2 widespread populations with measurable gene flow between them. Neutrality tests and mismatch distribution of the mitochondrial sequences indicated a departure from mutation-drift equilibrium due to a population expansion at the 2 extremes of the geographic range, but not towards the middle of the distribution. These results highlight the importance of evaluating multiple genetic markers to gain reliable insights into population processes and structure.

Indo-Pacific humpback dolphins (Sousa chinensis) in Hong Kong: Modelling demographic parameters with mark-recapture techniques.

Indo-Pacific humpback dolphins (Sousa chinensis) inhabiting Hong Kong waters are thought to be among the world's most anthropogenically impacted coastal delphinids. We have conducted a 5-year (2010-2014) photo-ID study and performed the first in this region comprehensive mark-recapture analysis applying a suite of open population models and robust design models. Cormack-Jolly-Seber (CJS) models suggested a significant transient effect and seasonal variation in apparent survival probabilities as result of a fluid movement beyond the study area. Given the spatial restrictions of our study, limited by an administrative border, if emigration was to be considered negligible the estimated survival rate of adults was 0.980. Super-population estimates indicated that at least 368 dolphins used Hong Kong waters as part of their range. Closed robust design models suggested an influx of dolphins from winter to summer and increased site fidelity in summer; and outflux, although less prominent, during summer-winter intervals. Abundance estimates in summer (N = 144-231) were higher than that in winter (N = 87-111), corresponding to the availability of prey resources which in Hong Kong waters peaks during summer months. We point out that the current population monitoring strategy used by the Hong Kong authorities is ill-suited for a timely detection of a population change and should be revised.

Threshold of long-term survival of a coastal delphinid in anthropogenically degraded environment: Indo-Pacific humpback dolphins in Pearl River Delta.

Defining demographic and ecological threshold of population persistence can assist in informing conservation management. We undertook such analyses for the Indo-Pacific humpback dolphin (Sousa chinensis) in the Pearl River Delta (PRD) region, southeast China. We use adult survival estimates for assessments of population status and annual rate of change. Our estimates indicate that, given a stationary population structure and minimal risk scenario, ~2000 individuals (minimum viable population in carrying capacity, MVPk) can maintain the population persistence across 40 generations. However, under the current population trend (~2.5% decline/annum), the population is fast approaching its viability threshold and may soon face effects of demographic stochasticity. The population demographic trajectory and the minimum area of critical habitat (MACH) that could prevent stochastic extinction are both highly sensitive to fluctuations in adult survival. For a hypothetical stationary population, MACH should approximate 3000-km2. However, this estimate increases four-fold with a 5% increase of adult mortality and exceeds the size of PRD when calculated for the current population status. On the other hand, cumulatively all current MPAs within PRD fail to secure the minimum habitat requirement to accommodate sufficiently viable population size. Our findings indicate that the PRD population is deemed to become extinct unless effective conservation measures can rapidly reverse the current population trend.

Spatiotemporal Trends of Heavy Metals in Indo-Pacific Humpback Dolphins (Sousa chinensis) from the Western Pearl River Estuary, China.

We assessed the spatiotemporal trends of the concentrations of 11 heavy metals (HMs) in the liver and kidney of Indo-Pacific humpback dolphins (Sousa chinensis) from western Pearl River Estuary (PRE) during 2004-2015. The hepatic levels of Cr, As, and Cu in these dolphins were among the highest reported for cetaceans globally, and the levels of Zn, Cu, and Hg were sufficiently high to cause toxicological effects in some of the animals. Between same age-sex groups, dolphins from Lingdingyang were significantly more contaminated with Hg, Se, and V than those from the West-four region, while the opposite was true for Cd. Generalized additive mixed models showed that most metals had significant but dissimilar temporal trends over a 10-year period. The concentrations of Cu and Zn increased significantly in recent years, corresponding to the high input of these metals in the region. Body-length-adjusted Cd levels peaked in 2012, accompanied by the highest annual number of dolphin stranding events. In contrast to the significant decrease in HM levels in the dolphins in Hong Kong waters (the eastern reaches of the PRE), the elevated metal exposure in the western PRE raises serious concerns.

Home range establishment and utilization by reintroduced lions (Panthera leo) in a small South African wildlife reserve.

Understanding of animal spatial behavior is essential for informed management decisions. In southern Africa, reintroduction of lions (Panthera leo) to small reserves (<1000 km2 ) has increased since the early 2000s, however studies on their ranging behavior in these enclosed systems remain lacking. We applied Time Local Convex Hull (T-LoCoH) methods to study the home range establishment and utilization of 11 lions reintroduced to Dinokeng Game Reserve, South Africa, during 2011 through 2014. Lions established home ranges close to their release sites and during the following 3 years their home range sizes continued to increase, but in each individual case the size remained smaller than half of the reserve area (<70 km2 ). Space use strategies differed between the core and the entire home range, with higher frequency of visits found in core areas. Exceptionally high rates (>60 separate visits) around the largest dam and along rivers suggest the importance of water and its surrounding vegetation in the lions' space utilization pattern. The home range size did not differ with season or sex of the individuals, whereas shifts in locations of home ranges revealed differences in the response of the 2 sexes to territorial conflicts and management interventions. Our study shows a dynamic home range utilization pattern and highlights the importance of both fine-scale space use patterns (frequency and duration of visits) and broad-scale home range changes in understanding the ranging behavior of reintroduced animals.

Increased human occupation and agricultural development accelerates the population contraction of an estuarine delphinid.

Over the past few thousand years, human development and population expansion in southern China have led to local extirpation and population contraction of many terrestrial animals. At what extent this early human-induced environmental change has also affected coastal marine species remains poorly known. We investigated the demographic history of the Indo-Pacific humpback dolphin (Sousa chinensis) in the Pearl River Delta (PRD); an obligatory inshore species known for its susceptibility to anthropogenic impacts in one of China's most developed coastal regions. Although the deltaic evolution of PRD has been influenced by climate since the Holocene, ~74% reduction of the dolphin's effective population size occurred within the last 2000 years, consistent with ~61% habitat contraction during this period. This considerable and recent population contraction may have been due to land use practices and deforestation in the upper/middle Pearl River region, all leading to increasing sedimentation rate in the estuarine area. As anthropogenic impacts within the drainage of Pearl River affected a vast area, coastal dolphins and large terrestrial mammals in southern China may share a similar demographic history, whilst the demographic and biogeographic history of the PRD humpback dolphins may be symptomatic of similar processes that this species may have undergone elsewhere in the region.

Profiling and Spatial Variation Analysis of Persistent Organic Pollutants in South African Delphinids.

The continuous disposal of persistent organic pollutants (POPs) in South Africa (SA) warrants concern about their detrimental effects on humans and wildlife. We surveyed six dolphin species (n = 90) incidentally captured in shark net installations or stranded off the SA east and south coast from 2005 to 2009 to study the POP exposure. Sousa plumbea, an inshore and estuarine species, was found to be the most contaminated by total POPs (21 100 ng g(-1) lw) of all the dolphins off SA, followed by Tursiops aduncus (19 800 ng g(-1) lw), Lagenodelphis hosei (13 600 ng g(-1) lw), and Delphinus capensis (5500 ng g(-1) lw), whereas POP levels in the offshore or pelagic delphinids were much lower. In all delphinids, dominant pollutants were dichlorodiphenyltrichloroethanes (DDTs), which represented more than 60% of the total concentration of total POPs, followed by polychlorinated biphenyls (PCBs, 30%). Concentrations of DDTs in S. plumbea and T. aduncus off SA were among the highest levels reported in delphinids globally. Approximately half of the adult T. aduncus had PCB concentrations above the effect threshold for impairment of immune functions. The concentrations of Mirex and Dieldrin in SA delphinids were higher than those found in species from other regions of the Southern Hemisphere.

Humpback Dolphins in Hong Kong and the Pearl River Delta: Status, Threats and Conservation Challenges.

In coastal waters of the Pearl River Delta (PRD) region, the Indo-Pacific humpback dolphin (Sousa chinensis) is thought to number approximately 2500 individuals. Given these figures, the putative PRD population may appear strong enough to resist demographic stochasticity and environmental pressures. However, living in close proximity to the world's busiest seaport/airport and several densely populated urban centres with major coastal infrastructural developments comes with challenges to the long-term survival of these animals. There are few other small cetacean populations that face the range and intensity of human-induced pressures as those present in the PRD and current protection measures are severely inadequate. Recent mark-recapture analyses of the animals in Hong Kong waters indicate that in the past two decades the population parameters have not been well understood, and spatial analyses show that only a very small proportion of the dolphins' key habitats are given any form of protection. All current marine protected areas within the PRD fail to meet a minimum habitat requirement that could facilitate the population's long-term persistence. Demographic models indicate a continuous decline of 2.5% per annum, a rate at which the population is likely to drop below the demographic threshold within two generations and lose 74% of the current numbers within the lifespan of three generations. In Hong Kong, the case of humpback dolphins represents a particularly explicit example of inadequate management where a complete revision of the fundamental approach to conservation management is urgently needed.

Phylogeography of the finless porpoise (genus Neophocaena): testing the stepwise divergence hypothesis in the northwestern Pacific.

We used 344 mitochondrial control region (717 bp) sequences from the finless porpoise (genus Neophocaena) from the northwestern Pacific to investigate the extent and manner in which past climatic oscillations may have shaped patterns of genetic diversity for this marine mammal. Both SplitsTree and Analysis of Molecular Variance (AMOVA) revealed the presence of a deep divergence among N. phocaenoides in subtropical waters compared with N. asiaeorientalis in temperate waters. Results from Migrate-n indicated that migration increased along the continent during the early Pleistocene period. Migration increased, although to a lesser extent than that during the Pleistocene, along the marginal shelf in the Yellow/Bohai Sea during the Last Glacial Maximum (LGM) due to a shortening coastline. Our results suggest that the current patterns of genetic diversity of Neophocaena vary at a hierarchy on a temporal and spatial scale, and phylogeographic history should be taken into account when examining species population structure and taxonomy.

Integrating multiple lines of evidence to better understand the evolutionary divergence of humpback dolphins along their entire distribution range: a new dolphin species in Australian waters?

The conservation of humpback dolphins, distributed in coastal waters of the Indo-West Pacific and eastern Atlantic Oceans, has been hindered by a lack of understanding about the number of species in the genus (Sousa) and their population structure. To address this issue, we present a combined analysis of genetic and morphologic data collected from beach-cast, remote-biopsied and museum specimens from throughout the known Sousa range. We extracted genetic sequence data from 235 samples from extant populations and explored the mitochondrial control region and four nuclear introns through phylogenetic, population-level and population aggregation frameworks. In addition, 180 cranial specimens from the same geographical regions allowed comparisons of 24 morphological characters through multivariate analyses. The genetic and morphological data showed significant and concordant patterns of geographical segregation, which are typical for the kind of demographic isolation displayed by species units, across the Sousa genus distribution range. Based on our combined genetic and morphological analyses, there is convincing evidence for at least four species within the genus (S. teuszii in the Atlantic off West Africa, S. plumbea in the central and western Indian Ocean, S. chinensis in the eastern Indian and West Pacific Oceans, and a new as-yet-unnamed species off northern Australia).

The evolving male: spinner dolphin (Stenella longirostris) ecotypes are divergent at Y chromosome but not mtDNA or autosomal markers.

The susceptibility of the Y chromosome to sexual selection may make this chromosome an important player in the formation of reproductive isolating barriers, and ultimately speciation. Here, we investigate the role of the Y chromosome in phenotypic divergence and reproductive isolation of spinner dolphin (Stenella longirostris) ecotypes. This species contains six known ecotypes (grouped into four subspecies) that exhibit striking differences in morphology, habitat and mating system, despite having adjacent or overlapping ranges and little genetic divergence at previously studied mtDNA and autosomal markers. We examined the phylogeographic structure for all six ecotypes across the species range (n = 261, 17 geographic locations) using DNA sequences from three Y chromosome markers, two maternally inherited mitochondrial (mtDNA) markers, and a biparentally inherited autosomal intron. mtDNA and autosomal analyses revealed low divergence (most Φ(ST) values <0.1) between ecotypes and geographic regions, concordant with previous studies. In contrast, Y intron analyses revealed fixed differences amongst the three most phenotypically divergent groups: S. l. longirostris vs. S. l. roseiventris vs. combined S. l. orientalis/S. l. centroamericana/Tres Marias ecotypes). Another ecotype (whitebelly), previously postulated to be a hybrid between the two phenotypically most divergent ecotypes, had Y haplotypes from both putative parent ecotypes, supporting a hybrid designation. Reduced introgression of the Y chromosome has previously been observed in other organisms ranging from insects to terrestrial mammals, and here we demonstrate this phenomenon in a marine mammal with high dispersal capabilities. These results indicate that reduced introgression of the Y chromosome occurs in a wide taxonomic range of organisms and support the growing body of evidence that rapid evolution of the Y chromosome is important in evolutionary diversification.

Permanent genetic resources added to Molecular Ecology Resources Database 1 February 2012 - 31 March 2012.

This article documents the addition of 171 microsatellite marker loci and 27 pairs of single nucleotide polymorphism (SNP) sequencing primers to the Molecular Ecology Resources Database. Loci were developed for the following species: Bombus pauloensis, Cephalorhynchus heavisidii, Cercospora sojina, Harpyhaliaetus coronatus, Hordeum vulgare, Lachnolaimus maximus, Oceanodroma monteiroi, Puccinia striiformis f. sp. tritici, Rhea americana, Salmo salar, Salmo trutta, Schistocephalus solidus, Sousa plumbea and Tursiops aduncus. These loci were cross-tested on the following species: Aquila heliaca, Bulweria bulwerii, Buteo buteo, Buteo swainsoni, Falco rusticolus, Haliaeetus albicilla, Halobaena caerulea, Hieraaetus fasciatus, Oceanodroma castro, Puccinia graminis f. sp. Tritici, Puccinia triticina, Rhea pennata and Schistocephalus pungitii. This article also documents the addition of 27 sequencing primer pairs for Puffinus baroli and Bulweria bulwerii and cross-testing of these loci in Oceanodroma castro, Pelagodroma marina, Pelecanoides georgicus, Pelecanoides urinatrix, Thalassarche chrysostoma and Thalassarche melanophrys.

Echolocation signals of Heaviside's dolphins (Cephalorhynchus heavisidii).

Field recordings of echolocation signals produced by Heaviside's dolphins (Cephalorhynchus heavisidii) were made off the coast of South Africa using a hydrophone array system. The system consisted of three hydrophones and an A-tag (miniature stereo acoustic data-logger). The mean centroid frequency was 125 kHz, with a -3 dB bandwidth of 15 kHz and -10 dB duration of 74 µs. The mean back-calculated apparent source level was 173 dB re 1 µPa(p.-p.). These characteristics are very similar to those found in other Cephalorhynchus species, and such narrow-band high-frequency echolocation clicks appear to be a defining characteristic of the Cephalorhynchus genus. Click bursts with very short inter-click intervals (up to 2 ms) were also recorded, which produced the "cry" sound reported in other Cephalorhynchus species. Since inter-click intervals correlated positively to click duration and negatively to bandwidth, Heaviside's dolphins may adjust their click duration and bandwidth based on detection range. The bimodal distribution of the peak frequency and stable bimodal peaks in spectra of individual click suggest a slight asymmetry in the click production mechanism.

Rolling stones and stable homes: social structure, habitat diversity and population genetics of the Hawaiian spinner dolphin (Stenella longirostris).

Spinner dolphins (Stenella longirostris) exhibit different social behaviours at two regions in the Hawaiian Archipelago: off the high volcanic islands in the SE archipelago they form dynamic groups with ever-changing membership, but in the low carbonate atolls in the NW archipelago they form long-term stable groups. To determine whether these environmental and social differences influence population genetic structure, we surveyed spinner dolphins throughout the Hawaiian Archipelago with mtDNA control region sequences and 10 microsatellite loci (n = 505). F-statistics, Bayesian cluster analyses, and assignment tests revealed population genetic separations between most islands, with less genetic structuring among the NW atolls than among the SE high islands. The populations with the most stable social structure (Midway and Kure Atolls) have the highest gene flow between populations (mtDNA Phi(ST) < 0.001, P = 0.357; microsatellite F(ST) = -0.001; P = 0.597), and a population with dynamic groups and fluid social structure (the Kona Coast of the island of Hawai'i) has the lowest gene flow (mtDNA 0.042 < Phi(ST) < 0.236, P < 0.05; microsatellite 0.016 < F(ST) < 0.040, P < 0.001). We suggest that gene flow, dispersal, and social structure are influenced by the availability of habitat and resources at each island. Genetic comparisons to a South Pacific location (n = 16) indicate that Hawaiian populations are genetically depauperate and isolated from other Pacific locations (mtDNA 0.216 < F(ST) < 0.643, P < 0.001; microsatellite 0.058 < F(ST) < 0.090, P < 0.001); this isolation may also influence social and genetic structure within Hawai'i. Our results illustrate that genetic and social structure are flexible traits that can vary between even closely-related populations.

The social structure and strategies of delphinids: predictions based on an ecological framework.

Dolphins live in complex social groupings with a wide variety of social strategies. In this chapter we investigate the role that differing habitats and ecological conditions have played in the evolution of delphinid social strategies. We propose a conceptual framework for understanding natural patterns of delphinid social structure in which the spatial and temporal predictability of resources influences the ranging patterns of individuals and communities. The framework predicts that when resources are spatially and temporally predictable, dolphins should remain resident in relatively small areas. Predictable resources are often found in complex inshore environments where dolphins may hide from predators or avoid areas with high predator density. Additionally, available food resources may limit group size. Thus, we predict that there are few benefits to forming large groups and potentially many benefits to being solitary or in small groups. Males may be able to sequester solitary females, controlling mating opportunities. Observations of inshore populations of bottlenose dolphins (Tursiops sp.) and island-associated spinner dolphins (Stenella longirostris) seem to fit this pattern well, along with forest-dwelling African antelope and primates such as vervets (Cercopithicus aethiops), baboons (Papio sp.), macaques (Macaca sp.) and chimpanzees (Pan troglodytes). In contrast, the framework predicts that when resources such as food are unpredictable, individuals must range further to find the necessary resources. Forming groups may be the only strategy available to avoid predation, especially in the open ocean. Larger home ranges are likely to support a greater number of individuals; however, prey is often sparsely distributed, which may act to reduce foraging competition. Cooperative foraging and herding of prey schools may be advantageous, potentially facilitating the formation of long-term bonds. Alternately, individuals may display many short-term affiliations. These large groups make it difficult for a male or a small group of males to sequester a female, and polygynandry is the most likely mating strategy. While it is difficult to study wide-ranging delphinids to examine these predictions, this ranging and behavioural pattern has been suggested for dusky dolphins (Lagenorhynchus obscurus), coastal bottlenose dolphins (Tursiops sp.) and mixed species of dolphins in the Eastern Tropical Pacific. These patterns also resemble the ranging and social strategies of open savannah African antelopes and desert-dwelling macropods. Resource availability exists in a range of complex distributions and we predict that delphinid ranging patterns will also vary. At intermediate-ranging patterns, the framework predicts that individuals should form mid-sized groups balancing intra-group competition with predation protection. Humpback dolphins (Sousa sp.) appear to fit this pattern, with some site fidelity over relatively large ranges. They display fluid associations with other individuals. Predation pressure is not sufficiently high to cause large groups to form, and individuals probably reduce predation pressure more by hiding whenever possible. This pattern is likely to prevent the formation of long-term complex bonds. In contrast, killer whales (Orcinus orca) also display intermediate-ranging patterns, but have extremely strong social bonds within familial groups. Cooperative and altruistic behaviour in killer whales facilitate the formation of life-long bonds, similar to those observations in sperm whales (Physeter macrocephalus) and elephants (Loxodonta africana). This conceptual framework remains largely untested, and for many species it is not currently possible to describe ranging behaviours, anti-predator tactics or social behaviour in sufficient detail for appropriate examination of these ideas. Few studies on dolphins have been conducted to explicitly test this type of framework; however, existing observations of delphinid social strategies and communities are used throughout this chapter to examine this framework. Additionally, we anticipate that the present framework may provide a starting point to test hypotheses regarding the evolution of social strategies of delphinids.