Low Genetic Variability in the Hawaiian Monk Seal

The Hawaiian monk seal (   Monachus schauinslandi) is a critically endangered species that has failed to recover from human exploitation despite decades of protection and ongoing management efforts designed to increase population growth. The seals breed at five principal locations in the northwestern Hawaiian islands, and inter-island migration is limited. Genetic variation in this species is expected to be low due to a recent population bottleneck and probable inbreeding within small subpopulations. To test the hypothesis that small population size and strong site fidelity has led to low within-island genetic variability and significant between-island differentiation, we used two independent approaches to quantify genetic variation both within and among the principal subpopulations. Mitochondrial control region and tRNA gene sequences (359 base pairs) were obtained from 50 seals and revealed very low genetic diversity (0.6% variable sites), with no evidence of subpopulation differentiation. Multilocus DNA fingerprints from 22 individuals also indicated low genetic variation in at least some subpopulations (band-sharing values for "unrelated" seals from the same island ranged from 49 to 73%). This method also provided preliminary evidence of population subdivision (  F'_st estimates of 0.20 and 0.13 for two adjacent island pairs). Translocations of seals among islands may therefore have the potential to relieve local inbreeding and possibly to reduce the total amount of variation preserved in the population. Genetic variation is only one of many factors that determine the ability of an endangered species to recover. Maintenance of existing genetic diversity, however, remains an important priority for conservation programs because of the possibility of increased disease resistance in more variable populations and the chance that inbreeding depression may only be manifest under adverse environmental conditions.     

Microsatellite Analysis of Population Structure in the Endangered Hawaiian Monk Seal

Abstract: The endangered Hawaiian monk seal breeds at six locations in the northwestern Hawaiian Islands. To determine whether significant genetic differentiation exists among these sites, we used microsatellite loci to examine the monk seal population structure at the five largest breeding colonies. Of 27 loci isolated from other seal species, only 3 were polymorphic in an initial screening of one individual from each breeding site. Only two alleles were found at each of these 3 loci in samples of 46–108 individuals. This extremely low variation is consistent with other measures of genetic variability in this species and is probably the result of a recent severe population bottleneck, combined with a long-term history of small population sizes. Although the smallest monk seal subpopulation in this study ( Kure Atoll) showed some evidence of heterozygote deficit, possibly indicative of inbreeding, the next smallest ( Pearl and Hermes Reef) had an apparent excess of heterozygous individuals. Genetic differentiation was detected between the two subpopulations at extreme ends of the range ( Kure and French Frigate Shoals). This trend was significant only at the microsatellite locus for which we had the largest sample size ( Hg6.3: R _ST = 0.206, p = 0.002; allelic goodness of fit G _h = 15.412, p < 0.005). French Frigate Shoals is the source population for translocated animals that have been released primarily at Kure Atoll. Differentiation between these sites consisted of allele frequency differences (with the same allele predominant in each location at all three loci), rather than the preservation of alternative alleles. Although the translocations have had positive demographic effects, we recommend continued genetic monitoring of both the source and recipient populations because translocated individuals are now entering the breeding population.     

Range‐Wide Genetic Connectivity of the Hawaiian Monk Seal and Implications for Translocation

Abstract: The Hawaiian monk seal (Monachus schauinslandi) is one of the most critically endangered marine mammals. Less than 1200 individuals remain, and the species is declining at a rate of approximately 4% per year as a result of juvenile starvation, shark predation, and entanglement in marine debris. Some of these problems may be alleviated by translocation; however, if island breeding aggregates are effectively isolated subpopulations, moving individuals may disrupt local adaptations. In these circumstances, managers must balance the pragmatic need of increasing survival with theoretical concerns about genetic viability. To assess range‐wide population structure of the Hawaiian monk seal, we examined an unprecedented, near‐complete genetic inventory of the species (n =1897 seals, sampled over 14 years) at 18 microsatellite loci. Genetic variation was not spatially partitioned (_w=?0.03, p = 1.0), and a Bayesian clustering method provided evidence of one panmictic population (K =1). Pairwise F_ST comparisons (among 7 island aggregates over 14 annual cohorts) did not reveal temporally stable, spatial reproductive isolation. Our results coupled with long‐term tag‐resight data confirm seal movement and gene flow throughout the Hawaiian Archipelago. Thus, human‐mediated translocation of seals among locations is not likely to result in genetic incompatibilities.     

Shifting Baselines and The Extinction of The Caribbean Monk Seal

The recent extnction of the Caribbean monk seal Monachus tropicalis has been considered an example of a human‐caused extinction in the marine environment, and this species was considered a driver of the changes that have occurred in the structure of Caribbean coral reef ecosystems since colonial times. I searched archaeological records, historical data, and geographic names (used as a proxy of the presence of seals) and evaluated the use and quality of these data to conclude that since prehistoric times the Caribbean monk seal was always rare and vulnerable to human predation. This finding supports the hypothesis that in AD 1500, the Caribbean monk seal persisted as a small fragmented population in which individuals were confined to small keys, banks, or isolated islands in the Gulf of Mexico and the Caribbean Sea. This hypothesis is contrary to the assumption that the species was widespread and abundant historically. The theory that the main driver of monk seal extinction was harvesting for its oil for use in the sugar cane industry of Jamaica during the 18th century is based primarily on anecdotal information and is overemphasized in the literature. An analysis of reported human encounters with this species indicates monk seal harvest was an occasional activity, rather than an ongoing enterprise. Nevertheless, given the rarity of this species and its restricted distribution, even small levels of hunting or specimen collecting must have contributed to its extinction, which was confirmed in the mid‐20th century. Some sources had been overlooked or only partially reviewed, others misinterpreted, and a considerable amount of anecdotal information had been uncritically used. Critical examination of archaeological and historical records is required to infer accurate estimations of the historical abundance of a species. In reconstructing the past to address the shifting baseline syndrome, it is important to avoid selecting evidence to confirm modern prejudices. Puntos de Referencia Cambiantes y la Extinción de la Foca Monje Caribeña     

Evolutionary Consequences of Extinctions in Populations of a Hawaiian Honeycreeper

We report on the evolutionary change in bill size of a species of Hawaiian honeycreeper resulting from an apparent dietary shift caused by dramatic declines and extinctions of lobelioids, a historically favored nectar source. Although it now feeds mainly on the flowers of the ohia tree ( Metrosideros polymorpha ), early Hawaiian avifaunal accounts report that the i'iwi ( Vestiaria coccinea ), which has a long decurved bill, fed primarily on the flowers of Hawaiian Lobelioideae, which typically have long decurved corollas. A coevolutionary association of i'iwi bill and flower morphology has often been asserted. We test the hypothesis that the shift in the i'iwi's diet from the long corolla lobelioid flowers to ohia flowers, which lack corollas, resulted in directional selection for shorter bills. We evaluate this hypothesis by comparing the morphological characters of museum specimens from the island of Hawaii collected before 1902 with recent specimens from the Hakalau National Wildlife Refuge, Hawaii. We examine evidence of change in morphological characters using multivariate analysis and a nonparametric cubic spline technique. Results from all analyses are congruent: bill length is shorter in recent specimens.     

Life and Death in the Fast Lane: Demographic Consequences of Road Mortality in the Florida Scrub-Jay

Abstract: We examined the demographic consequences of road mortality in the cooperatively breeding Florida Scrub-Jay (Aphelocoma coerulescens ), a threatened species restricted to the oak scrub of peninsular Florida. Between May 1986 and July 1995 we monitored the survival and reproductive success of a color-banded population of jays along a two-lane highway at Archbold Biological Station. Annual mortality of breeding adults was 0.38 on road territories, significantly higher than the rate of 0.23 for breeders on nonroad territories. High mortality on road territories appeared to be a direct result of automobile traffic per se and not a consequence of road-induced changes in habitat characteristics. Mortality was especially high for immigrants without previous experience living along the road: in their first two years as breeders on road territories, naive immigrants experienced annual mortality of 0.50 and 0.45. From year 3 onward, however, annual mortality dropped to 0.29, not significantly different from the rate for birds on nonroad territories. This experience-dependent decline in road mortality could be caused either by surviving jays learning to avoid automobiles or by selective mortality operating through time (demographic heterogeneity). Proximity to the road had no effect on nesting success beyond its indirect effects on breeder experience and group size. Because the mortality of 30- to 90-day-old fledglings was significantly higher on road territories than on nonroad territories, however, breeder mortality greatly exceeded production of yearlings on road territories. Roadside territories therefore are sinks that can maintain populations of Florida Scrub-Jays only via immigration. Because Florida Scrub-Jays do not avoid roadside habitats and may even be attracted to them, road mortality presents a difficult challenge for the management and conservation of this threatened and declining species.     

"Mobbing" in Hawaiian Monk Seals (Monachus schauinslani): The Value of Simulation Modeling in the Absence of Apparently Crucial Data

Several small populations of Hawaiian monk seals ( Monachus schauinslani ) exhibit male-biased adult sex ratios and "mobbing," an aggressive behavior in which adult males injure and often kill adult females and immature seals of both sexes during mating attempts. Mobbing appears to be limiting the growth of some populations of this endangered species. The frequency of mobbing deaths appears to increase as a population's sex ratio becomes increasingly male-biased, although the exact relationship between these two variables (the mobbing response) is unknown. We developed a stochastic demographic model of a small Hawaiian monk seal population using several different assumptions about the mobbing response. We used the model to explore the origins of male-biased sex ratios in monk seal populations and to determine whether it was possible, given the lack of data on the mobbing response, to evaluate the probable effects of alternative management strategies to address the mobbing problem. Small populations (100 to 200 seals) and those with slower growth rates were more likely to develop male-biased adult sex ratios. Almost all of our modeling scenarios supported the immediate removal of males from populations where mobbing occurs. Our conclusions were relatively unaffected when the assumptions regarding the mobbing response were varied. Thus, a model was helpful even when apparently crucial data were unavailable.     

Introduced Birds and the Fate of Hawaiian Rainforests

Abstract:  The Hawaiian Islands have lost nearly all their native seed dispersers, but have gained many frugivorous birds and fleshy-fruited plants through introductions. Introduced birds may not only aid invasions of exotic plants but also may be the sole dispersers of native plants. We assessed seed dispersal at the ecotone between native- and exotic-dominated forests and quantified bird diets, seed rain from defecated seeds, and plant distributions. Introduced birds were the primary dispersers of native seeds into exotic-dominated forests, which may have enabled six native understory plant species to become reestablished. Some native plant species are now as common in exotic forest understory as they are in native forest. Introduced birds also dispersed seeds of two exotic plants into native forest, but dispersal was localized or establishment minimal. Seed rain of bird-dispersed seeds was extensive in both forests, totaling 724 seeds of 9 native species and 2 exotics with over 85% of the seeds coming from native plants. Without suitable native dispersers, most common understory plants in Hawaiian rainforests now depend on introduced birds for dispersal, and these introduced species may actually facilitate perpetuation, and perhaps in some cases restoration, of native forests. We emphasize, however, that restoration of native forests by seed dispersal from introduced birds, as seen in this study, depends on the existence of native forests to provide a source of seeds and protection from the effects of ungulates. Our results further suggest that aggressive control of patches of non-native plants within otherwise native-dominated forests may be an important and effective conservation strategy.     

Demographic Consequences of Terrestrial Habitat Loss for Pool‐Breeding Amphibians: Predicting Extinction Risks Associated with Inadequate Size of Buffer Zones

Abstract: Much of the biodiversity associated with isolated wetlands requires aquatic and terrestrial habitat to maintain viable populations. Current federal wetland regulations in the United States do not protect isolated wetlands or extend protection to surrounding terrestrial habitat. Consequently, some land managers, city planners, and policy makers at the state and local levels are making an effort to protect these wetland and neighboring upland habitats. Balancing human land‐use and habitat conservation is challenging, and well‐informed land‐use policy is hindered by a lack of knowledge of the specific risks of varying amounts of habitat loss. Using projections of wood frog (Rana sylvatica) and spotted salamander (Ambystoma maculatum) populations, we related the amount of high‐quality terrestrial habitat surrounding isolated wetlands to the decline and risk of extinction of local amphibian populations. These simulations showed that current state‐level wetland regulations protecting 30 m or less of surrounding terrestrial habitat are inadequate to support viable populations of pool‐breeding amphibians. We also found that species with different life‐history strategies responded differently to the loss and degradation of terrestrial habitat. The wood frog, with a short life span and high fecundity, was most sensitive to habitat loss and isolation, whereas the longer‐lived spotted salamander with lower fecundity was most sensitive to habitat degradation that lowered adult survival rates. Our model results demonstrate that a high probability of local amphibian population persistence requires sufficient terrestrial habitat, the maintenance of habitat quality, and connectivity among local populations. Our results emphasize the essential role of adequate terrestrial habitat to the maintenance of wetland biodiversity and ecosystem function and offer a means of quantifying the risks associated with terrestrial habitat loss and degradation.     

Habitat and reef-fish assemblages of banks in the Northwestern Hawaiian Islands

The moderately deep terraces and banks of the Northwestern Hawaiian Islands (NWHI) were surveyed to describe their habitat and reef-fish assemblages. These tracts of bottom at 30–40 m comprise more than 4,500 km² of the regions reef area. The habitat was found to be dominated by algal meadows (65% cover of exposed bottom), with infrequent relief features. Annual monitoring of select stations for 4 years at Necker Bank indicated that the relative difference in algal abundance between stations persisted from year to year (at least in summer). Temperature records from year-long deployments of archival thermistors in high-cover (>70%) and low-cover (<30%) algal biotopes were indistinguishable, providing no explanation of the algal differences between stations. At all banks, Microdictyon was the primary alga, averaging 1.22 kg/m². In spite of the extensive standing primary production, and a historical lack of fishing, bank reef-fish populations were impoverished. Mean densities, sizes, and biomass of trophic groups were considerably less than values reported for NWHI reef shallows. An overall mean biomass was estimated at 22.5 g/m², which is a fifth of that reported for shallow reefs of the region. Fish biomass of all trophic groups was associated with the few sources of relief available on the banks. Apex predators (sharks, jacks, and snappers), common on all surveys (with a mean of five per station), were proposed to constrain fish populations to sparse sources of relief resulting in a skewed size structure of the two primary fish trophic components. Sizes of lower-level carnivores were tightly correlated with sources of relief whereas the size of herbivores were not, indicating that herbivores more often venture out and risk the exposed algal meadows. These bank summits are a rare example of a near pristine reef system with high benthic primary productivity and low fish biomass, and are a stark contrast to shallower coral-reef ecosystems of the NWHI.Communicated by P.W. Sammarco, Chauvin     

Effects of temperature on the mortality and growth of Hawaiian reef corals

Three common species of Hawaiian reef corals, Pocillopora damicornis (L.), Montipora verrucosa (Lamarck) and Fungia scutaria Lamarck, were grown in a temperature-regulated, continuous-flow sea water system. The skeletal growth optimum occurred near 26°C, coinciding with the natural summer ambient temperature in Hawaii, and was lowest at 21° to 22°C, representing Hawaiian winter ambient. Levels of approximately 32°C produced mortality within days. Prolonged exposure to temperatures of approximately 30°C eventually caused loss of photosynthetic pigment, increased mortality, and reduced calcification. Corals lived only 1 to 2 weeks at 18°C. The corals showed greater initial resistance at the lower lethal limit, but ultimately low temperature was more deleterious than high temperature. Results suggest that a decrease in the natural water temperature of Hawaiian reefs would be more harmful to corals than a temperature increase of the same magnitude.Contribution No. 504 of the Hawaii Institute of Marine Biology.     

Endemism and dispersal: comparative phylogeography of three surgeonfishes across the Hawaiian Archipelago

To evaluate the hypothesis that a general correlation exists between species range size and dispersal ability, we surveyed mitochondrial cytochrome b sequence variation in three surgeonfish species with vastly different ranges: Ctenochaetus strigosus, Hawaiian endemic, N = 531; Zebrasoma flavescens, North Pacific, N = 560; Acanthurus nigrofuscus, Indo-Pacific, N = 305. Collections were made throughout the 2,500 km expanse of the Hawaiian Archipelago and adjacent Johnston Atoll. Analyses of molecular variance demonstrate that all three species are capable of maintaining population connectivity on a scale of thousands of km (all species global Φ_ST = NS). However, rank order comparison of pairwise Φ_ST results and Exact test P-values revealed modest but significantly different patterns of gene flow among the three species surveyed, with the degree of genetic structure increasing as range size decreases (P = 0.001). These results are consistent with mtDNA surveys of four additional Hawaiian reef fauna in which a wide-spread Indo-Pacific species exhibited genetic homogeneity across the archipelago, while three endemics had significant population subdivision over the same range. Taken together, these seven cases invoke the hypothesis that Hawaii’s endemic reef fishes evolved from species with reduced dispersal ability that, after initial colonization, could not maintain contact with parent populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Securing the Demographic and Genetic Future of Tuatara through Assisted Colonization

Abstract: Climate change poses a particular threat to species with fragmented distributions and little or no capacity to migrate. Assisted colonization, moving species into regions where they have not previously occurred, aims to establish populations where they are expected to survive as climatic envelopes shift. However, adaptation to the source environment may affect whether species successfully establish in new regions. Assisted colonization has spurred debate among conservation biologists and ecologists over whether the potential benefits to the threatened species outweigh the potential disruption to recipient communities. In our opinion, the debate has been distracted by controversial examples, rather than cases where assisted colonization may be a viable strategy. We present a strategic plan for the assisted migration of tuatara (Sphenodon punctatus), an endemic New Zealand reptile. The plan includes use of extant populations as reference points for comparisons with assisted‐colonization populations with respect to demography, phenotypic plasticity, and phenology; optimization of genetic variation; research to fill knowledge gaps; consideration of host and recipient communities; and inclusion of stakeholders in the planning stage. When strategically planned and monitored, assisted colonization could meet conservation and research goals and ultimately result in the establishment of long‐term sustainable populations capable of persisting during rapid changes in climate.     

Size structure of phytoplankton biomass and photosynthesis in subtropical Hawaiian waters

In a subtropical Hawaiian ecosystem, phytoplanton size structure analyses (November–December, 1980) showed that ultraplankton (>3μm), nanoplankton (>20μm) and netplankton (>20μm) accounted for ca. 80, 98, and 2% of total chlorophyll standing stock, respectively, on the basis of chlorophyll. Similar trends were evident, for other biomass indices (e.g. cell numbers, total cell volume, ATP, particulate organic carbon, particulate organic nitrogen). The ultraplankton fraction consisted primarily of small flagellates (1 to 3 μm diam) and coccoid cells (?1 μm diam); the 3 to 20 μm fraction was represented by dinoflagellates, coccolithophores, diatoms, and chrysophytes; and the netplankton fraction consisted principally of dinoflagellates and centric diatoms. Community photosynthesis had a size distribution similar to that of biomass. Sinking rates for the 3 μm, 3 to 20 μm, and >20 μm fractions averaged 0.0, 0.09, and 0.29m d^?1, respectively. The absence of measurable sinking rates for the ultraplankton, together with the relative abundance of biomass in this fraction, result in very small phytoplankton losses due to sinking in such subtropical surface waters.     

Annual,semilunar, and diel reproductive rhythms in the Hawaiian labrid Thalassoma duperrey

In September 1978, 5 034 juvenile western rock (spiny) lobsters (Panulirus cygnus George) were tagged on a shallow coastal reef near Cliff Head in Western Australia. Of these, 26% were recaptured and removed from the population during the subsequent commercial fishery between 15 November 1978 and 30 June 1979. A resurvey of the tagging site in September 1979 found that size structure of the population was similar to the tagged population in September 1978 but that the 1979 population was composed of almost completely different individuals and only 2% of the rock lobsters tagged in September 1978 were recaught at this time. A wide range of movement was indicated by the recapture positions of 354 rock lobsters caught off the tagging site. Most travelled almost directly offshore but some also travelled offshore in a northwesterly direction. Both trends were shown in the data from the recaptures of 18 rock lobster released farther offshore from Cliff Head in September 1981, and of 19 released at a close inshore site in earlier years. Maximum offshore rates of movement of up to 622 m d^-1 were recorded for distances up to 37 nautical miles (ca. 68 km). Growth rates for the tagged individuals which moved off, and those that were caught on, the tagging site were similar and close to growth rate under optimal conditions for this species. Under the present high levels of exploitation few, if any, of the rock lobsters remain to become permanent residents in the coastal reefs. It is believed that the north-westerly movements of some of the pre-adult rock lobsters, on their way to the breeding grounds on the outer edge of the continental shelf, are due to the upstream movement of these rock lobsters in a current from the north. The north-westerly movements allow the rock lobster population increased utilization of the food supplies of the coastal area.     

Risk Analysis of Hunting of Seal Populations in the Baltic

Vulnerabilities of grey seal ( Halichoerus grypus) and ringed seal ( Phoca hispida) populations in the Baltic Sea were evaluated for potential opening of the populations for hunting. We used ecological risk analysis to assess the effects of environmental and demographic stochasticity and uncertain and partly missing population data on the modeled outcomes. The impact of different harvesting strategies on the long-term sustainability of seal populations was analyzed with four different models with increasing complexity and population detail. It appears the simpler the population model used, the more overconfident results it gave with regard to the hunting policy to be adopted. Therefore, it proves risky in population management decisions to rely on simplistic calculations based on growth rate and estimated population size alone. This is even more so if the population estimates have a wide error margin. Due to an unknown, but presumably positive number of seal kills in the Baltic at present, the sustainable harvest is likely to be close to zero for both seal species. Our risk analysis strongly suggests refraining from Baltic seal hunting now, with their current population sizes, and in the future if the development of population numbers cannot be assessed accurately enough.     

Genetic and Demographic Threats to the Black Rhinoceros Population in the Ngorongoro Crater

A resident population of 13 black rhinoceros ( Diceros bicornis ) persist in Ngorongoro Crater, Tanzania. The effective population size ( N _e ) may be as few as 5 animals. Projected growth for this population suggests that the effective population size will remain small for the near future, threatening this Iocal population with extinction due to the stochastic factors associated with small population size. A summary of historic and recent demographic data for this population reveals a population crash during the period of heavy poaching that affected this species throughout its range. Although poaching of this species has been brought under control the population remains small. These data and models of projected population growth argue for consideration of more-intensive management within the framework of the small population paradigm. This case is an example of applied conservation resulting from this paradigm used in conjunction with rather than competing with the declining population paradigm. We identify additional monitoring, particularly of density-dependent behaviors, that will be necessary for designing a successful management program. Finally, the use of molecular markers for developing an accurate pedigree for this population is suggested in order to maintain a genetically healthy population. These strategies have broad applicability to black rhinoceros conservation throughout Africa.     

Demographic Side Effects of Selective Hunting in Ungulates and Carnivores

Abstract:  Selective harvesting regimes are often implemented because age and sex classes contribute differently to population dynamics and hunters show preferences associated with body size and trophy value. We reviewed the literature on how such cropping regimes affect the demography of the remaining population (here termed demographic side effects ). First, we examined the implications of removing a large proportion of a specific age or sex class. Such harvesting strategies often bias the population sex ratio toward females and reduce the mean age of males, which may consequently delay birth dates, reduce birth synchrony, delay body mass development, and alter offspring sex ratios. Second, we reviewed the side effects associated with the selective removal of relatively few specific individuals, often large trophy males. Such selective harvesting can destabilize social structures and the dominance hierarchy and may cause loss of social knowledge, sexually selected infanticide, habitat changes among reproductive females, and changes in offspring sex ratio. A common feature of many of the reported mechanisms is that they ultimately depress recruitment and in some extreme cases even cause total reproductive collapse. These effects could act additively and destabilize the dynamics of populations, thus having a stronger effect on population growth rate than first anticipated. Although more experimental than observational studies reported demographic side effects, we argue that this may reflect the quite subtle mechanisms involved, which are unlikely to be detected in observational studies without rigorous monitoring regimes. We call for more detailed studies of hunted populations with marked individuals that address how the expression of these effects varies across mating systems, habitats, and with population density. Theoretical models investigating how strongly these effects influence population growth rates are also required.