Predictive Modeling of Spinner Dolphin (Stenella longirostris) Resting Habitat in the Main Hawaiian Islands

Predictive habitat models can provide critical information that is necessary in many conservation applications. Using Maximum Entropy modeling, we characterized habitat relationships and generated spatial predictions of spinner dolphin (Stenella longirostris) resting habitat in the main Hawaiian Islands. Spinner dolphins in Hawai'i exhibit predictable daily movements, using inshore bays as resting habitat during daylight hours and foraging in offshore waters at night. There are growing concerns regarding the effects of human activities on spinner dolphins resting in coastal areas. However, the environmental factors that define suitable resting habitat remain unclear and must be assessed and quantified in order to properly address interactions between humans and spinner dolphins. We used a series of dolphin sightings from recent surveys in the main Hawaiian Islands and a suite of environmental variables hypothesized as being important to resting habitat to model spinner dolphin resting habitat. The model performed well in predicting resting habitat and indicated that proximity to deep water foraging areas, depth, the proportion of bays with shallow depths, and rugosity were important predictors of spinner dolphin habitat. Predicted locations of suitable spinner dolphin resting habitat provided in this study indicate areas where future survey efforts should be focused and highlight potential areas of conflict with human activities. This study provides an example of a presence-only habitat model used to inform the management of a species for which patterns of habitat availability are poorly understood.     

Site fidelity and association patterns in a deep-water dolphin: Rough-toothed dolphins (Steno bredanensis) in the Hawaiian Archipelago

In the Pacific, rough-toothed dolphins ( Steno bredanensis ) are typically found in the open ocean and in deep waters around oceanic islands. We examined habitat use, site fidelity, movements, and association patterns of this species in the main Hawaiian Islands. Sighting rates were highest in depths >1,500 m. There were frequent within- and between-year resightings off the island of Hawai'i, indicating a small population size with high site fidelity. Resighting rates were lower off Kaua'i/Ni'ihau, indicating a larger population size, but with some site fidelity. Two individuals were documented moving from Kaua'i to Hawai'i, a distance of 480 km, but were not seen to associate with dolphins off Hawai'i. Observed movements were consistent with at most 2% dispersal per year between these two areas. Differences in group sizes, habitat use, and behavior imply that movements among the islands may be limited. Little is known about the diet of rough-toothed dolphins in Hawai'i, but they are thought to feed primarily on near-surface species. High fidelity to deep-water areas off the island of Hawai'i likely reflects an increase in the predictability of prey associated with upwelling due to the island mass effect, wind stress curl and cyclonic eddies that form off the island.     

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 Φ_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 < Φ_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.     

Population structure of island‐associated dolphins: Evidence from mitochondrial and microsatellite markers for common bottlenose dolphins (Tursiops truncatus) around the main Hawaiian Islands

We used mitochondrial and nuclear genetic markers to investigate population structure of common bottlenose dolphins, Tursiops truncatus, around the main Hawaiian Islands. Though broadly distributed throughout the world's oceans, bottlenose dolphins are known to form small populations in coastal waters. Recent photo‐identification data suggest the same is true in Hawaiian waters. We found genetic differentiation among (mtDNA Φ_ST= 0.014–0.141, microsatellite F’_ST= 0.019–0.050) and low dispersal rates between (0.17–5.77 dispersers per generation) the main Hawaiian Island groups. Our results are consistent with movement rates estimated from photo‐identification data and suggest that each island group supports a demographically independent population. Inclusion in our analyses of samples collected near Palmyra Atoll provided evidence that the Hawaiian Islands are also occasionally visited by members of a genetically distinct, pelagic population. Two of our samples exhibited evidence of partial ancestry from Indo‐Pacific bottlenose dolphins (T. aduncus), a species not known to inhabit the Hawaiian Archipelago. Our findings have important implications for the management of Hawaiian bottlenose dolphins and raise concerns about the vulnerability to human impacts of pelagic species in island ecosystems.     

SPINNER DOLPHINS (STENELLA LONGIROSTRIS) OF THE WESTERN PACIFIC AND SOUTHEAST ASIA: PELAGIC AND SHALLOW-WATER FORMS1

A dwarf form of the spinner dolphin has been reported from the Gulf of Thailand, while more typical large spinner dolphins have been described from Japanese waters and other localities in the western Pacific. These reports have been based on very few specimens. Our purpose in this study was to determine the affinities of spinner dolphins throughout the region based on larger samples and to review their taxonomic status, with an hypothesis of two widespread ecotypic forms, or subspecies. We examined 213 osteological specimens, from a tuna gillnet fishery in the Philippines, from the former Taiwanese shark gillnet fishery in the Timor and Arafura Seas off northern Australia, from the Gulf of Thailand, from other areas in the western Pacific and Southeast Asia, from the eastern Indian Ocean, and from the Central and South Pacific. Results show that spinner dolphins from the deep inner waters of the Philippines conform to the large pelagic type of spinner dolphin that inhabits the Central and South Pacific, the western Pacific and the eastern Indian Ocean. The skull is similar in size and shape to the holotype specimen of S. longirostris (from unknown locality). This form feeds primarily on small mesopelagic fishes and squids. Spinner dolphins from the shallow waters of inner Southeast Asia represented in the sample, including the Gulf of Thailand, Timor Sea and Arafura Sea, are smaller in body and skull size, have fewer teeth and vertebrae, and feed mainly on benthic and coral reef fishes and invertebrates. We hypothesize that this form also inhabits the Java Sea and other shallow waters throughout inner Indonesia and Malaysia. We redescribe a subspecies corresponding to the small form and based on Delphinus roseiventris Wagner 1846 from the Arafura Sea, designating a neotype and paraneotype specimens.     

Differences in oscillatory whistles produced by spinner (Stenella longirostris) and pantropical spotted (Stenella attenuata) dolphins

Acoustic recordings of two closely related species, spinner dolphin (Stenella longirostris) and pantropical spotted dolphin (Stenella attenuata), were investigated from four different geographic locations: two in the Central Tropical Pacific, one in the Eastern Tropical Pacific and one in the Indian Ocean. The two delphinid species occur in tropical and warm temperate waters, with overlapping ranges. They produce very similar vocalizations, but at the same time their calls exhibit a certain degree of intraspecific variation among different geographic locations as has been observed in other delphinid species. Oscillatory whistles (whistles with at least two oscillations in their frequency contours) were identified and manually extracted from the recordings. Whistles with four or more maxima (oscillations) occurred only in spinner dolphins and they were present in all geographic regions investigated. In addition, the oscillatory whistles with two and three maxima were significantly more frequent in spinner than in spotted dolphins. The differences in oscillatory whistles for these two species seem to be consistent across study areas and therefore, could be used in addition to other whistle features to help distinguish between them.     

Genetic differentiation in endemic Silene (Caryophyllaceae) on the Hawaiian Islands

Endemic Hawaiian Silene colonize new lava flows and are outcompeted as the ground ages. They illustrate the genetic processes operating in the evolution of pioneering island plants. The volcanic history of the Hawaiian Islands allows an estimation of the age of these plant populations. In this study, populations of S. struthioloides from Maui and the older part of the island of Hawaii, and S. hawaiiensis from the youngest volcanoes of the island of Hawaii were analyzed by enzyme electrophoresis. The genetic structures of these populations were placed in a geographic and geologic context. Silene is much more polymorphic on the older island, Maui. Genetic variation appears to have been lost in the colonization of the youngest island, Hawaii. Interestingly, some loci for populations on the younger volcanoes of the island of Hawaii are monomorphic for alleles not found in populations on the older part of Hawaii. Recurrent colonizations are accompanied with founder effects, and restricted gene flow among populations has led to isolation and genetic drift. This has resulted in genetic and morphological differentiâtion of Silene populations on the youngest volcanoes.     

Tracing ancestry with chromosomal sequences

Most of the large Drosophila species of Hawaii are single-island endemics. Chromosomal sequences show that species at the new end of the archipelago have been derived stepwise from ancestral populations on older islands. The oldest high island has an endemic species with sequences that match some in the Nearctic-Palearctic robusta species group. Colonization from a continent by long-distance dispersal seems a likely origin for the Hawaiian drosophilids. Telmatogeton, a worldwide genus of marine midges, has five Hawaiian species inhabiting freshwater streams. Chromosomal sequences of a marine species in Hawaiian waters match the freshwater forms, indicating colonization from the ocean.     

Mid-ocean isolation and the evolution of Hawaiian reef fishes

The Hawaiian fish fauna has close affinities with the fauna of the Indo-west Pacific from which it is derived, but is depauperate. It is characterized by a large number of endemic species (30% of inshore fishes), which are often the most abundant species in their families in Hawaii. Although there is evidence of local adaptation, there has been no radiation of species within the island chain, as occurs in the terrestrial biota of isolated islands. Three major factors have contributed to these trends: (1) the geographic isolation of the islands, and oceanographic features, especially current patterns; (2) the life history characteristics of the fishes, especially their dispersal capabilities; and (3) the extent of adaptive differentiation to environmental conditions after they reached Hawaii.     

Population structure of island-associated dolphins: Evidence from photo-identification of common bottlenose dolphins (Tursiops truncatus) in the main Hawaiian Islands

Management agencies often use geopolitical boundaries as proxies for biological boundaries. In Hawaiian waters a single stock is recognized of common bottlenose dolphins, Tursiops truncatus , a species that is found both in open water and near-shore among the main Hawaiian Islands. To assess population structure, we photo-identified 336 distinctive individuals from the main Hawaiian Islands, from 2000 to 2006. Their generally shallow-water distribution, and numerous within-year and between-year resightings within island areas suggest that individuals are resident to the islands, rather than part of an offshore population moving through the area. Comparisons of identifications obtained from Kaua'i/Ni'ihau, O'ahu, the "4-island area," and the island of Hawai'i showed no evidence of movements among these island groups, although movements from Kaua'i to Ni'ihau and among the "4-islands" were documented. A Bayesian analysis examining the probability of missing movements among island groups, given our sample sizes for different areas, indicates that interisland movement rates are less than 1% per year with 95% probability. Our results suggest the existence of multiple demographically independent populations of island-associated common bottlenose dolphins around the main Hawaiian islands.     

New evidence of ectomycorrhizal fungi in the Hawaiian Islands associated with the endemic host Pisonia sandwicensis (Nyctaginaceae)

Ectomycorrhizal plants and fungi are ubiquitous in mainland forests, but because of dispersal limitations are predicted to be less common on isolated islands. For instance, no native ectomycorrhizal plants or fungi have ever been reported from Hawaii, one of the most remote archipelagos on Earth. Members of the plant tribe Pisonieae are common on many islands, and prior evidence shows that some species associate with ectomycorrhizal fungi. However, until now, the Pisonieae species of Hawaii had yet to be examined for their mycorrhizal status. Here we sampled roots from members of the genus Pisonia growing on the Hawaiian islands of Oahu, Maui and Hawaii. We used molecular and microscopic techniques to categorize trees with respect to their mycorrhizal associations. We report that the Hawaiian endemic Pisonia sandwicensis forms ectomycorrhizas with at least five fungal operational taxonomic units (corresponding closely to species) belonging to four genera. We also report that this tree species is monophyletic with other ectomycorrhizal Pisonia species. We suggest that in light of the newly discovered Hawaiian ectomycorrhizal fungal community and other island ectomycorrhizal communities, dispersal limitations do not prevent the colonization of remote islands by at least some ectomycorrhizal fungi.     

Darwin Point: A threshold for atoll formation

Summary A threshold for atoll formation, herein termed the Darwin Point, exists at the northern end of the Hawaiian Archipelago at 29°N latitude. Hawaiian atolls and coral islands transported northwest by tectonic movement of the Pacific Plate appear to have drowned near the Darwin Point during the last 20 million years. Measures of gross carbonate production by corals across the archipelago show that growth rates decrease with increasing latitude. At the Darwin Point, corals may contribute only 20% of the calcium carbonate necessary to keep pace with recent changes in sea level and thus appear to be more important as builders of framework than producers of limestone. Reduction in this function rather than total carbonate production may be the determining factor in the formation of atolls and coral islands. Elsewhere in the world other Darwin Points may exist but probably not at the same latitude due to differences in ecological conditions, coral species composition, island area, rates of erosion and tectonic histories.Hawaii Institute of Marine Biology Contribution No. 627     

Phylogeographic patterns of Hawaiian Megalagrion damselflies (Odonata: Coenagrionidae) correlate with Pleistocene island boundaries

The Pleistocene geological history of the Hawaiian Islands is becoming well understood. Numerous predictions about the influence of this history on the genetic diversity of Hawaiian organisms have been made, including the idea that changing sea levels would lead to the genetic differentiation of populations isolated on individual volcanoes during high sea stands. Here, we analyse DNA sequence data from two closely related, endemic Hawaiian damselfly species in order to test these predictions, and generate novel insights into the effects of Pleistocene glaciation and climate change on island organisms. Megalagrion xanthomelas and Megalagrion pacificum are currently restricted to five islands, including three islands of the Maui Nui super-island complex (Molokai, Lanai, and Maui) that were connected during periods of Pleistocene glaciation, and Hawaii island, which has never been subdivided. Maui Nui and Hawaii are effectively a controlled, natural experiment on the genetic effects of Pleistocene sea level change. We confirm well-defined morphological species boundaries using data from the nuclear EF-1alpha gene and show that the species are reciprocally monophyletic. We perform phylogeographic analyses of 663 base pairs (bp) of cytochrome oxidase subunit II (COII) gene sequence data from 157 individuals representing 25 populations. Our results point to the importance of Pleistocene land bridges and historical island habitat availability in maintaining inter-island gene flow. We also propose that repeated bottlenecks on Maui Nui caused by sea level change and restricted habitat availability are likely responsible for low genetic diversity there. An island analogue to northern genetic purity and southern diversity is proposed, whereby islands with little suitable habitat exhibit genetic purity while islands with more exhibit genetic diversity.     

Stepping stone speciation in Hawaii’s flycatchers: molecular divergence supports new island endemics within the elepaio

The elepaio (Chasiempis sandwichensis) is a monarch flycatcher endemic to the Hawaiian Islands of Kauai, Oahu, and Hawaii. Elepaio vary in morphology among and within islands, and five subspecies are currently recognized. We investigated phylogeography of elepaio using mitochondrial (ND2) and nuclear (LDH) markers and population structure within Hawaii using ND2 and microsatellites. Phylogenetic analyses revealed elepaio on each island formed reciprocally monophyletic groups, with Kauai ancestral to other elepaio. Sequence divergence in ND2 among islands (3.02–2.21%) was similar to that in other avian sibling species. Estimation of divergence times using relaxed molecular clock models indicated elepaio colonized Kauai 2.33 million years ago (95% CI 0.92–3.87 myr), Oahu 0.69 (0.29–1.19) myr ago, and Hawaii 0.49 (0.21–0.84) myr ago. LDH showed less variation than ND2 and was not phylogenetically informative. Analysis of molecular variance within Hawaii showed structure at ND2 (fixation index = 0.31), but microsatellites showed no population structure. Genetic, morphological, and behavioral evidence supports splitting elepaio into three species, one on each island, but does not support recognition of subspecies within Hawaii or other islands. Morphological variation in elepaio has evolved at small geographic scales within islands due to short dispersal distances and steep climatic gradients. Divergence has been limited by lack of dispersal barriers in the extensive forest that once covered each island, but anthropogenic habitat fragmentation and declines in elepaio population size are likely to decrease gene flow and accelerate differentiation, especially on Oahu.     

Bats are able to maintain long-term social relationships despite the high fission-fusion dynamics of their groups

Elephants, dolphins, as well as some carnivores and primates maintain social links despite their frequent splitting and merging in groups of variable composition, a phenomenon known as fission-fusion. Information on the dynamics of social links and interactions among individuals is of high importance to the understanding of the evolution of animal sociality, including that of humans. However, detailed long-term data on such dynamics in wild mammals with fully known demography and kin structures are scarce. Applying a weighted network analysis on 20,500 individual roosting observations over 5 years, we show that in two wild Bechstein's bat colonies with high fission-fusion dynamics, individuals of different age, size, reproductive status and relatedness maintain long-term social relationships. In the larger colony, we detected two stable subunits, each comprising bats from several family lineages. Links between these subunits were mainly maintained by older bats and persisted over all years. Moreover, we show that the full details of the social structure become apparent only when large datasets are used. The stable multi-level social structures in Bechstein's bat colonies resemble that of elephants, dolphins and some primates. Our findings thus may shed new light on the link between social complexity and social cognition in mammals.     

Population genetic structure and taxonomy of the common dolphin (Delphinus sp.) at its southernmost range limit: New Zealand waters

New Zealand is the southernmost limit of the common dolphin's (genus Delphinus) distribution in the Pacific Ocean. In this area, common dolphins occur in both coastal and oceanic habitats, exhibit seasonal and resident occurrence, and present high morphological variability. Here we investigated the population structure and the taxonomic identity of common dolphins (Delphinus sp.) within New Zealand waters using 14 microsatellite loci, 577 bp of the mtDNA control region, and 1,120 bp of the mtDNA cytochrome b gene across 90 individuals. We found high genetic variability and evidence of population expansion. Phylogenetic analyses conducted to clarify the taxonomic status of New Zealand common dolphins did not show any clustering reflecting geographic origin or morphotypes. The microsatellite analysis showed genetic differentiation between Coastal and Oceanic putative populations, while mtDNA revealed significant genetic differentiation only between the Hauraki Gulf and other putative groups. Our results suggest that differences in habitat choice and possible female site fidelity may play a role in shaping population structure of New Zealand common dolphins.     

Between a rock and a hard place: habitat selection in female-calf humpback whale (Megaptera novaeangliae) Pairs on the Hawaiian breeding grounds

The Au'au Channel between the islands of Maui and Lanai, Hawaii comprises critical breeding habitat for humpback whales (Megaptera novaeangliae) of the Central North Pacific stock. However, like many regions where marine mega-fauna gather, these waters are also the focus of a flourishing local eco-tourism and whale watching industry. Our aim was to establish current trends in habitat preference in female-calf humpback whale pairs within this region, focusing specifically on the busy, eastern portions of the channel. We used an equally-spaced zigzag transect survey design, compiled our results in a GIS model to identify spatial trends and calculated Neu's Indices to quantify levels of habitat use. Our study revealed that while mysticete female-calf pairs on breeding grounds typically favor shallow, inshore waters, female-calf pairs in the Au'au Channel avoided shallow waters (<20 m) and regions within 2 km of the shoreline. Preferred regions for female-calf pairs comprised water depths between 40-60 m, regions of rugged bottom topography and regions that lay between 4 and 6 km from a small boat harbor (Lahaina Harbor) that fell within the study area. In contrast to other humpback whale breeding grounds, there was only minimal evidence of typical patterns of stratification or segregation according to group composition. A review of habitat use by maternal females across Hawaiian waters indicates that maternal habitat choice varies between localities within the Hawaiian Islands, suggesting that maternal females alter their use of habitat according to locally varying pressures. This ability to respond to varying environments may be the key that allows wildlife species to persist in regions where human activity and critical habitat overlap.     

A DWARF FORM OF THE SPINNER DOLPHIN (STENELLA LONGIROSTRIS) FROM THAILAND

A very small form of the spinner dolphin has been found to inhabit the Gulf of Thailand. Ten specimens taken incidentally in a local shrimp fishery differ from specimens of this species collected elsewhere in body size and shape, skull size and shape, number of teeth and numbers of thoracic and lumbar vertebrae. Four cranially adult males were 129–137 cm long, well below the ranges for the Indian Ocean, western Pacific, central Pacific, eastern Pacific and Atlantic. The skull is also very small. Tooth counts and vertebral counts average lower than in other series. The color pattern is not significantly different from that of spinner dolphins in the central and western Pacific, Atlantic and Indian Oceans but differs from that of the small eastern spinner of the eastern Pacific. The Gulf of Thailand specimens are morphologically separable from all other specimens, but it is to be expected that when larger samples are available there will be some overlap. The dwarf form may overlap in body and skull size with small spinner dolphins taken incidentally in a gillnet fishery for sharks off northern Australia. The geographic range of the dwarf spinner may be restricted. The size and status of the population and the impact of the shrimp fishery are unknown and should be investigated. The dwarf spinner may have an ecology different from that of other spinners, feeding mainly on reef-associated and benthic organisms rather than mesopelagic animals.     

Site fidelity,residency, and abundance of bottlenose dolphins (Tursiops sp.) in Adelaide's coastal waters,South Australia

Little is known about the ecology and behavior of southern Australian bottlenose dolphins (Tursiops sp.). This hinders assessment of their conservation status and informed decision‐making concerning their management. We used boat‐based surveys and photo‐identification data to investigate site fidelity, residency patterns, and the abundance of southern Australian bottlenose dolphins in Adelaide's coastal waters. Sighting rates and site fidelity varied amongst individuals, and agglomerative hierarchical cluster analysis led to the categorization of individuals into one of three groups: occasional visitors, seasonal residents, or year‐round residents. Lagged identification rates indicated that these dolphins used the study area regularly from year to year following a model of emigration and reimmigration. Abundance estimates obtained from multisample closed capture‐recapture models ranged from 95 individuals (SE ± 45.20) in winter 2013 to 239 (SE ± 54.91) in summer 2014. The varying levels of site fidelity and residency, and the relatively high number of dolphins found throughout the study area highlights the Adelaide metropolitan coast as an important habitat for bottlenose dolphins. As these dolphins also appear to spend considerable time outside the study area, future research, conservation, and management efforts on this population must take into account anthropogenic activities within Adelaide's coastal waters and their adjacencies.     

Restricted dispersal in a continuously distributed marine species: common bottlenose dolphins Tursiops truncatus in coastal waters of the western North Atlantic

The marine environment provides an opportunity to examine population structure in species with high dispersal capabilities and often no obvious barriers to genetic exchange. In coastal waters of the western North Atlantic, common bottlenose dolphins, Tursiops truncatus, are a highly mobile species with a continuous distribution from New York to Florida. We examine if the highly mobile nature coupled with no obvious geographic barriers to movement in this region result in a large panmictic population. Mitochondrial control region sequences and 18 microsatellite loci indicate dolphins are partitioning the habitat both latitudinally and longitudinally. A minimum of five genetically differentiated populations were identified among 404 samples collected in the range of New Jersey to northern Florida using both genetic marker types, some inhabiting nearshore coastal waters and others utilizing inshore estuarine waters. The genetic results reject the hypothesis of a single stock of coastal bottlenose dolphins put forth after the 1987–1988 epizootic that caused a large‐scale die‐off of dolphins and suggest instead the disease vector was transferred from one population to the next as a result of seasonal migratory movements of some populations. These coastal Atlantic populations also differ significantly from bottlenose dolphin samples collected in coastal waters of the northern Gulf of Mexico, implying a long‐term barrier to movement between the two basins.