Ranging patterns of bottlenose dolphins living in oceanic waters: implications for population structure

Very little is known about the ecology of common bottlenose dolphins (Tursiops truncatus) living in oceanic waters. This study investigated the ranging and residence patterns of bottlenose dolphins occurring in the Azores (Portugal), the most isolated archipelago in the North Atlantic. Data were collected during standardized boat-based surveys conducted over a 6-year period in an area of approximately 5,400 km² (main study area). To investigate the extent of movements of individual animals, non-systematic surveys were also conducted outside this area. Only 44 individuals out of 966 identified were frequently sighted within and between years. The remaining individuals were either temporary migrants from within or outside the archipelago, or transients. Resident dolphins showed strong geographic fidelity to the area. Long-distance movements (of almost 300 km), consistent with foraging or exploratory trips, were observed among non-resident dolphins. Home range size was estimated for 31 individuals sighted ≥10 times. Range areas of these dolphins varied in size and location, but considerable overlap was observed in the areas used, suggesting the absence of habitat partitioning between resident and non-resident dolphins. Estimates of home range size of bottlenose dolphins in the Azores were found to be considerably larger than those previously reported for this species. It is hypothesized that dolphins living in the Azores carry out extensive movements and have large home ranges in response to the lower density and patchy distribution of prey compared to other areas. The extensive ranging behaviour and the lack of territoriality provide an opportunity for interbreeding between dolphins associated with different islands, thus preventing genetic differentiation within the population of the Azores.     

Ranging patterns of bottlenose dolphins living in oceanic waters: implications for population structure

Very little is known about the ecology of common bottlenose dolphins (Tursiops truncatus) living in oceanic waters. This study investigated the ranging and residence patterns of bottlenose dolphins occurring in the Azores (Portugal), the most isolated archipelago in the North Atlantic. Data were collected during standardized boat-based surveys conducted over a 6-year period in an area of approximately 5,400 km² (main study area). To investigate the extent of movements of individual animals, non-systematic surveys were also conducted outside this area. Only 44 individuals out of 966 identified were frequently sighted within and between years. The remaining individuals were either temporary migrants from within or outside the archipelago, or transients. Resident dolphins showed strong geographic fidelity to the area. Long-distance movements (of almost 300 km), consistent with foraging or exploratory trips, were observed among non-resident dolphins. Home range size was estimated for 31 individuals sighted ≥10 times. Range areas of these dolphins varied in size and location, but considerable overlap was observed in the areas used, suggesting the absence of habitat partitioning between resident and non-resident dolphins. Estimates of home range size of bottlenose dolphins in the Azores were found to be considerably larger than those previously reported for this species. It is hypothesized that dolphins living in the Azores carry out extensive movements and have large home ranges in response to the lower density and patchy distribution of prey compared to other areas. The extensive ranging behaviour and the lack of territoriality provide an opportunity for interbreeding between dolphins associated with different islands, thus preventing genetic differentiation within the population of the Azores.     

Investigating stock structure and trophic relationships among island-associated dolphins in the oceanic waters of the North Atlantic using fatty acid and stable isotope analyses

Short-beaked common dolphins (Delphinus delphis) and Atlantic spotted dolphins (Stenella frontalis) are the two most abundant cetacean species in the oceanic waters of Madeira and the Azores. They are of similar size, occur in similar habitats and are regularly observed in mixed-species groups to forage together. Genetic analyses suggested that, within each species, dolphins ranging around both archipelagos belong to the same panmictic population. We tested the hypotheses that (1) within each species, individuals from the two archipelagos belong to a single ecological stock; (2) between species, common and spotted dolphins have distinct trophic niches; using fatty acid (FA) and stable isotope (SI) analyses. Fatty acids and stable isotopes were analysed from 86 blubber and 150 skin samples of free-ranging dolphins, respectively. Sex-related differences were not significant, except for common dolphin FA profiles. In S. frontalis, FA and SI differences between archipelagos suggested that individuals belonged to different ecological stocks, despite the existence of gene flow between the two archipelagos. In D. delphis, differences were more pronounced, but it was not possible to distinguish between stock structure and a seasonal effect, due to differential sampling periods in the Azores and Madeira. Inter-specific comparisons were restricted to the Azores where all samples were collected during summer. Differences in FA proportions, noticeably for FA of dietary origin, as well as in nitrogen SI profiles, confirmed that both species feed on distinct resources. This study emphasizes the need for an integrated approach including both genetic and biochemical analyses for stock assessment, especially in wide-ranging marine top predators.     

The population structure of the harbour porpoise,Phocoena phocoena,in Danish waters and part of the North Atlantic

A study of the homogeneity between putative stocks of Phocoena phocoena in the North Sea and inner Danish waters was performed using isozyme electrophoresis. Two polymorphic systems, mannose phosphate isomerase (MPI) and phosphoglucomutase (PGM) were used in the analysis. A two-locus homogeneity test showed an effect of season and locality on the genotypic distribution. This led to a division of the total sample into season and into the two localities, the inner Danish waters (IDW) and the North Sea. The samples were then compared to the Hardy-Weinberg expectations where a deficit in heterozygotes was observed in the North Sea sample in both systems and at the Pgm locus in the total sample, indicating a mixing of sub-populations. A hierarchical contingency table analysis, also based on the genotypic distribution, showed a combined effect of season and locality and only an insignificant effect of sampling period, thus supporting the division into season and locatily. The seasonal effect is further supported by earlier observations of a seasonal migration of harbour porpoises out of the Baltic in winter. A significant difference was observed between the two supposed sub-populations, IDW-summer and North Sea-summer samples, based on the genotypic distribution. This could also be a reflection of the sample sizes. On the basis of samples obtained from Canada, West Greenland and Holland a preliminary study of the population structure on a larger scale, comprising the North East and North West Atlantic, was performed. The results so far accord well with the hitherto accepted sub-populations of P. phocoena in the North Atlantic.     

Distribution of common dolphins (Delphinus spp.) in the western Atlantic Ocean: a critical re-examination

Due to indications that misidentification (largely confusion among dolphins of the genera Delphinus and Stenella) in the past had led to erroneous assumptions of distribution of the two species of common dolphins (Delphinus delphis and D. capensis) in the western Atlantic Ocean, we conducted a critical re-examination of records of the genus Delphinus from this region. We compiled 460 ‘plottable’ records, required support for confirmation of genus and species identifications, and found many records lacking (and some clearly misidentified). When we plotted only the valid records (n = 364), we found evidence of populations in only three areas, and apparent absence throughout much of the tropical/subtropical regions. Off the east coast of the US and Canada, D. delphis is found from the Georgia/South Carolina border (32°N) north to about 47–50°N off Newfoundland. Since the 1960s, they have apparently been absent from Florida waters. There is no evidence that dolphins of the genus occur in the Gulf of Mexico. Reports of common dolphins from most of the Caribbean Basin are also rejected, and the only place in that region where they are confirmed to occur is off central-eastern Venezuela (a coastal D. capensis population). Off eastern South America, common dolphins appear to be restricted to south of 20°S. There is a coastal long-beaked population found in the South Brazil Bight, and one or more short-beaked populations south and offshore of this (ranging south to at least northern Argentina). The results are very different from commonly-accepted patterns of distribution for the genus in the Atlantic. Most areas of distribution coincide with moderate to strong upwelling and common dolphins appear to avoid warm, tropical waters. This study shows that great care must be taken in identification of similar-appearing long-beaked delphinids, and that uncritical acceptance of records at face value can lead to incorrect assumptions about the ranges of the species involved.     

Vertical community structure and ontogenetic distribution of chaetognaths in upper pelagic waters of the Eastern Mediterranean

The vertical distribution of chaetognaths and their developmental stages were investigated in the pelagic waters (0 to 500 m) of the Eastern Mediterranean Sea. Twenty vertical hauls were taken during autumn 1991 in the Ionian, Cretan, Levantine and Rhodes Seas. The chaetognaths Sagitta bipunctata and S. enflata had the same median depth distributions, which differed from those of S. minima, S. serratodentata, S. lyra, Krohnitta subtilis, and S. decipiens, while S. hexaptera occurred only sporadically. S. bipunctata and S. enflata were mainly restricted to <50 m, and while their distributions co-occurred their stage composition differed. The remaining species took up progressively deeper positions in the water column. Ontogenetic vertical distributions, with the older stages occurring at greater depth, were only observed for the mesopelagic species S. lyra and S. decipiens. Diurnal vertical migration was not detected in any of the species nor in their developmental stages.     

Phylogeography of two Atlantic squirrelfishes (Family Holocentridae): exploring links between pelagic larval duration and population connectivity

Genetic surveys of reef fishes have revealed high population connectivity within ocean basins, consistent with the assumption that pelagic larvae disperse long distances by oceanic currents. However, several recent studies have demonstrated that larval retention and self-recruitment may be higher than previously expected. To assess connectivity in tropical reef fishes, we contribute range-wide mtDNA surveys of two Atlantic squirrelfishes (family Holocentridae). The blackbar soldierfish, Myripristis jacobus, has a pelagic juvenile phase of about 58 days, compared to about 71 days (~22% longer) in the longjaw squirrelfish, Holocentrus ascensionis. If the pelagic duration is guiding dispersal ability, M. jacobus should have greater population genetic structure than H. ascensionis. In comparisons of mtDNA cytochrome b sequences from 69 M. jacobus (744 bp) and 101 H. ascensionis (769 bp), both species exhibited a large number of closely related haplotypes (h=0.781 and 0.974, π=0.003 and 0.006, respectively), indicating late Pleistocene coalescence of mtDNA lineages. Contrary to the prediction based on pelagic duration, M. jacobus has much less population structure (φ_ST=0.008, P=0.228) than H. ascensionis (φ_ST=0.091, P<0.001). Significant population partitions in H. ascensionis were observed between eastern, central and western Atlantic, and between Brazil and the Caribbean in the western Atlantic. These results, in combination with the findings from 13 codistributed species, indicate that pelagic larval duration is a poor predictor of population genetic structure in Atlantic reef fishes. A key to understanding this disparity may be the evolutionary depth among corresponding taxonomic groups of “reef fishes”, which extends back to the mid-Cretaceous and encompasses enormous diversity in ecology and life history. We should not expect a simple relationship between pelagic larval duration and genetic connectivity, among lineages that diverged 50–100 million years ago.     

Natural tracers in dietary studies: data for 210Po and 210Pb in decapod shrimp and other pelagic organisms in the Northeast Atlantic Ocean

Measurements of the concentrations of the naturally-occurring radioactive nuclides ²¹⁰Po and ²¹⁰Pb in a large number of pelagic organisms from the Atlantic Ocean made in the spring/summer of 1984 and 1985 are reported. Marine shrimp are particularly well represented, and in many cases data were obtained for shrimp hepatopancreas, stomach-contents, and posterior intestine plus contents, as well as for whole individuals. The wide ranges found in ²¹⁰Po concentrations group into categories: the shrimp, for example, divide into four categories, two penaeid and two carid. These groupings are explained on the basis of variations in diet between the different categories. A clear difference was also observed between the ²¹⁰Po levels in shrimp of similar species and size from opposite sides of an oceanographic front; this difference too can be explained plausibly in terms of a change in certain penaeid shrimp, and attention is drawn to the need for investigation of cytological and/or genetic effects which could possibly be caused by the very high natural radiation doses to which such organisms are exposed.     

On mathematical simulation of a pelagic ecosystem in tropical waters of the ocean

Consideration is given to a finite-difference model of functioning of a pelagic community inhabiting the upper 0 to 200 m layer of the tropical waters of the ocean. The model was developed on the basis of original data, obtained mainly on the 44th cruise of the R.V. Vityaz in the western Equatorial Pacific Ocean. Changes in the system are studied in time (up to 100 days, with 1 day intervals) and in depth (from surface to 200 m depth, with 10 m intervals). The state of the system in the upwelling zone, assumed to be characterized by a homogeneous vertical distribution of all its elements, is regarded as initial. The system then develops with time while it moves with the water flow. The model was realized on a BESM-3M computor. The computations yielded a pattern of changes in time of the biomass of the elements (expressed in calories), and a pattern of their vertical distribution at different instants. The vertical distribution pattern of the major elements of the model shows a rather close correlation with their observed distribution in the ocean at the corresponding instant.     

Comparative population genetic structure of marine gastropods (Littorina spp.) with and without pelagic larval dispersal

 Population genetic theory predicts that marine animal species with planktonic larvae will have less genetic structure than those with direct development. We compared the genetic structure of four species of littorinid snails – two with planktonic egg capsules that hatch as planktonic larvae and two with benthic egg masses that hatch as crawl-away juveniles. We used DNA sequencing and single stranded conformational polymorphism (SSCP) to assess sequence variation in a 480 bp fragment of the mitochondrial cytochrome b gene and then used an analysis of molecular variance (AMOVA) to estimate Φ_st for populations from the northeastern Pacific coast. One of the two direct-developing species, Littorina subrotundata, had a moderate amount of population structure (Φ_st=0.209) as expected but the other direct-developing species, L. sitkana, was nearly fixed for a single haplotype that made it impossible to precisely estimate Φ_st. One of the two planktonic-developing species, L. scutulata, did not show any significant population structure (Φ_st=0.004). In contrast to our expectations, the other planktonic-developing species, L. plena, showed some weak but statistically significant population structure (Φ_st=0.052). We discuss how differences in population genetic structure between species with the same type of development may reflect differences in their historical demography. Received: 22 December 1999 / Accepted: 24 July 2000     

Hydrology influences population genetic structure and connectivity of the intertidal amphipod Corophium volutator in the northwest Atlantic

The mechanisms driving genetic structure in marine systems are elusive due to the difficulty of identifying temporal and spatial barriers to dispersal. By studying marine invertebrate species with limited dispersal potential, genetic structure can be directly related to physical and biological factors restricting connectivity. In the northwest Atlantic, the benthic brood-rearing amphipod Corophium volutator is distributed across basins with distinct circulation patterns and has the potential to disperse passively during its adult stage. We analyzed spatial genetic variation and migration rates across C. volutator’s North American range with sequence data for mitochondrial DNA and three novel nuclear markers using frequency and coalescent-based methods. We found low genetic differentiation within basins, but strong subdivision within the Bay of Fundy and a striking biogeographic break between the Bay of Fundy and Gulf of Maine, suggesting that genetic drift may act on populations in which connectivity is restricted due to the limitation of passive dispersal by hydrological patterns.     

The relationship between population genetic structure and pelagic larval duration in coral reef fishes on the Great Barrier Reef

Pelagic larval duration (PLD) is a commonly used proxy for dispersal potential in coral reef fishes. Here we examine the relationship between PLD, genetic structure and genetic variability in geographically widespread and ecological generalist species from one coral reef fish family (Pomacentridae) that differs in mean larval duration by more than a month. The genetic structure was estimated in eight species using a mitochondrial molecular marker (D-loop) and in a sub-set of five species using nuclear molecular markers (ISSRs). Estimates of genetic differentiation were similar among species with pelagic larvae, but differed between molecular markers. The mtDNA indicated no structure in all species except one, while the ISSR indicated some structure between the sampling locations in all species. We detected a relationship between PLD and genetic structure using both markers. These relationships, however, were caused by a single species, Acanthochromis polyacanthus, which differs from all the other species examined here in lacking a larval phase. With this species excluded, there was no relationship between PLD and genetic structure using either marker despite a range of PLDs of more than 20 days. Genetic diversities were generally high in all species and did not differ significantly among species and locations. Nucleotide diversity and total heterozygosity were negatively related to maximum PLD but again these relationships were caused by A. polyacanthus and disappeared when this species was excluded. These genetic patterns are consistent with moderate gene flow among well-connected locations and indicate that at this phylogenetic level (i.e., within family) the duration of the pelagic larval phase is unrelated to the patterns of genetic differentiation.     

Population structure of short-beaked common dolphins (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) in the North Atlantic Ocean as revealed by mitochondrial and nuclear genetic markers

The understanding of population structure and gene flow of marine pelagic species is paramount to monitoring, management and conservation studies. Such studies are often hampered by the potentially high dispersal behavior of the species, the lack of obvious geographical barriers in the marine environment and the scarce sample availability. Short-beaked common dolphins (Delphinus delphis) are widespread in coastal and open-ocean habitats of the North Atlantic Ocean, nevertheless population structure and migratory patterns are poorly understood. Furthermore, concern has been raised about the status of the species because large numbers of dolphins have been taken incidentally in several fisheries throughout the North Atlantic in the past decades. In the present study, a large number of individual samples were obtained from seasonal and spatial aggregations of common dolphins from western (wNA) and eastern North Atlantic (eNA) regions, mostly using opportunistic sampling (i.e. from incidental entanglement in fishing gear or beach-cast carcasses). Genetic variability was investigated using nuclear (14 microsatellite loci) and mitochondrial (360 bp of the control region) genetic markers. Levels of genetic diversity were relatively high in all sampled areas and no evidence of recent reduction of effective population size (i.e. bottleneck) was detected at the nuclear loci. Significant population structure was detected between the two main regions (wNA and eNA) where it appeared to be more pronounced at mitochondrial (F _ST = 0.018, P < 0.001) than nuclear markers (F _ST = 0.005, P < 0.05), indicating the presence of at least two genetically distinct populations of common dolphins in the North Atlantic Ocean. In contrast, no significant genetic structure was detected between temporal aggregations of dolphins from within the same region, suggesting possible seasonal movement patterns at a regional scale. The observed levels of genetic differentiation between classes of markers are discussed here as a possible consequence of migratory patterns or recent population subdivision. An erratum to this article can be found at     

Phycoerythrin and photosynthesis of the pelagic blue-green alga Trichodesmium thiebautii in the waters of Kuroshio,Japan

The photosynthetic function of phycoerythrin was investigated in the alga Trichodesmium thiebautii from the waters of Kuroshio, Japan. The spectroscopic characteristics of the in vivo and isolated T. thiebautii phycoerythrin pigments are identical, and have 3 absorption bands at 495, 547 and 562 nm. Light at the wavelengths corresponding to each absorption band of phycoerythrin is equally efficient in T. thiebautii photosynthesis, indicating that phycoerythrin is active in trapping light energy for photosynthesis. In the natural habitat, phycoerythrin is considered to be the main photosynthetic pigment in T. thiebautii photosynthesis.     

Molecular phylogeny and population structure of tideland snails in the genus Cerithidea around Japan

Phylogenetic relationships and genetic population structures were analyzed for tideland gastropods in the genus Cerithidea around Japan on the basis of partial sequence of the mitochondrial COI gene. Large genetic divergence was shown between individuals of Cerithidea cingulata in the southern Ryukyus and those in the central Ryukyus and the Japanese Islands. Haplotypes of C. cingulata from the Japanese Islands were paraphyletic with the exclusion of a monophyletic group from the central Ryukyus. Genetic differentiation of C. cingulata was also detected between Amami-Oshima Island and Okinawajima Island. No genetic divergence was found between Cerithidea rhizophorarum in the Japanese Islands and its subspecies C. rhizophorarum morchii in the Ryukyu Islands. The lack of genetic divergence of Cerithidea largillierti between continental China and Japan suggests relatively recent migration between the Japanese Islands and the Asian continent. For all three Cerithidea species distributed in both the Japanese Islands and the Ryukyu Islands, the Tokara Gap and the Kerama Gap were shown to have acted as barriers to the dispersal.     

Population differentiation of the Atlantic spotted dolphin (Stenella frontalis) in the western North Atlantic, including the Gulf of Mexico

Information about the genetic population structure of the Atlantic spotted dolphin [Stenella frontalis (G. Cuvier 1829)] in the western North Atlantic would greatly improve conservation and management of this species in USA waters. To this end, mitochondrial control region sequences and five nuclear microsatellite loci were used to test for genetic differentiation of Atlantic spotted dolphins in the western North Atlantic, including the Gulf of Mexico (n=199). Skin tissue samples were collected from 1994–2000. Significant heterozygote deficiencies in three microsatellite loci within samples collected off the eastern USA coast prompted investigation of a possible Wahlund effect, resulting in evidence for previously unsuspected population subdivision in this region. In subsequent analyses including three putative populations, two in the western North Atlantic (n=38, n=85) and one in the Gulf of Mexico (n=76), significant genetic differentiation was detected for both nuclear DNA (R _ST=0.096, P≤0.0001) and mitochondrial DNA (Φ _ST=0.215, P≤0.0001), as well as for all pair-wise population comparisons for both markers. This genetic evidence for population differentiation coupled to known biogeographic transition zones at Cape Hatteras, North Carolina and Cape Canaveral, Florida, USA, evidence of female philopatry, and preliminary support for significant genetic differences between previously documented morphotypes of Atlantic spotted dolphins in coastal and offshore waters all indicate that the biology and life history of this species is more complex than previously assumed. Assumptions of large, panmictic populations might not be accurate in other areas where S. frontalis is continuously distributed (e.g., eastern Atlantic), and could have a detrimental effect on long-term viability and maintenance of genetic diversity in this species in regions where incidental human-induced mortality occurs.

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Mitochondrial DNA and electronic tracking reveal population structure of Atlantic bluefin tuna (Thunnus thynnus)

Population subdivision was examined in Atlantic bluefin tuna (Thunnus thynnus) through sequencing of the control region of the mitochondrial genome. A total of 178 samples from the spawning grounds in the Gulf of Mexico, Bahamas and Mediterranean Sea were analyzed. Among the samples from these locations were 36 electronically tagged bluefin tuna that were tagged in the North Atlantic and subsequently traveled to one of these known spawning grounds during the spawning season. Bluefin tuna populations from the Gulf of Mexico and the Mediterranean Sea were found to be genetically distinct based on Φ_st, and sequence nearest neighbor analyses, showing that these two major spawning areas support independent stocks. Sequence nearest neighbor analysis indicated significant population subdivision among the Gulf of Mexico, western Mediterranean and eastern Mediterranean Sea. However, it was not possible to find significant pairwise differences between any sampling areas when using all samples. If only samples that had a high likelihood of assignment to a specific spawning site were used (young of the year, spawning adults), the differentiation increased among all sampling areas and the Western Mediterranean Sea was distinct from the Eastern Mediterranean Sea and the Gulf of Mexico. It was not possible to distinguish samples from the Bahamas from those collected at any of the other sampling sites. These data support tagging results that suggested distinctness of the Gulf of Mexico, Eastern and Western Mediterranean Sea spawning areas. This level of stock differentiation is only possible if Atlantic bluefin tuna show strong natal homing to individual spawning grounds.     

Variation in pelagic larval growth of Atlantic billfishes: the role of prey composition and selective mortality

Atlantic blue marlin (Makaira nigricans) and sailfish (Istiophorus platypterus) larvae were collected from 10 monthly cruises (June–October 2003 and 2004) across the Straits of Florida to test (1) whether growth differed between the more productive western region near the Florida shelf, and the less productive eastern region toward the Bahamas, and (2) whether growth was related to prey consumption. Examination of larval sagittal otoliths revealed that instantaneous growth and daily growth during the first 2–3 weeks of life did not vary significantly between the two regions for either species. However, recent growth during the last two full days prior to collection was greater in the west for blue marlin larvae. Recent growth of blue marlin larvae <9 mm SL (primarily zooplanktivorous) was significantly related to prey composition (faster growth when higher proportions of Farranula copepods were consumed). Western larvae grew faster and had higher proportions of Farranula in their guts. Trends for sailfish larvae were not significant. In both species, comparison of early growth between <9 and ≥9 mm SL size groups indicated that growth trajectories diverged around 5–8 mm SL, the time when billfish larvae become capable of piscivory. Significantly faster growth of larger (older) larvae suggests that mortality was selective for fast growers and that the transition to piscivory may be a critical point in the early life of billfish.     

Fine-scale population structure of estuarine bottlenose dolphins (Tursiops truncatus) assessed using stable isotope ratios and fatty acid signature analyses

Stable isotope ratios and fatty acid signature analyses were employed to examine the fine-scale population structure of a year-round resident population of 600–800 bottlenose dolphins (Tursiops truncatus) in the Indian River Lagoon (IRL), Florida. The IRL, a 250-km-long estuary running along the central east coast of Florida (28.0°N, 80.6°W), is comprised of the northern and southern IRL, Mosquito Lagoon (ML), Banana River (BR), and St. Lucie Estuary. Samples of skin and blubber were collected from dead stranded (n = 61, 1994–2004) and live dolphins (n = 153, 2002–2007, 2010, 2011) from throughout the IRL and surrounding environs. Using stable isotopes (SI), dolphins could be assigned to a ML subpopulation, a St. Lucie Estuary subpopulation, and an IRL subpopulation. Fatty acid signature analysis (FASA) allowed for finer resolution, detecting ML and BR subpopulations, a separation of northern and southern IRL subpopulations, and a St. Lucie Estuary subpopulation. Differences between sexes were detectable within subpopulations using FASA, but not using SI. This may indicate that males and females are foraging in similar locations at a similar trophic level (detected using SI), but are varying in the types or proportions of specific prey (indicated by FASA). The combination of these complementary analyses results in a powerful tool for assessing fine-scale population substructure.