Sighting patterns reveal unobserved pupping events to revise reproductive rate estimates for Hawaiian monk seals in the main Hawaiian Islands

We used sighting reports, including decades of citizen-reported Hawaiian monk seal (Neomonachus schauinslandi) sightings, to describe female breeding biology and reproductive success in the main Hawaiian Islands. We first used this data set to describe the timing of events in the female reproductive cycle. We then conducted an expert review of patterns in sighting histories to detect unobserved pupping events. Finally, we estimated the age-specific reproductive curve for female monk seals in the main Hawaiian Islands. Charting reproductive cycles showed indications of the robust condition of female monk seals in the main Hawaiian Islands; they nursed pups 12% longer than their counterparts in the Northwestern Hawaiian Islands and regained condition to molt more quickly after weaning a pup. By examining sighting histories, we were able to infer 25 unobserved pupping events that had previously gone uncounted. We accounted for additional uncertainty with a randomization procedure. After accounting for unobserved pupping events, the age-specific reproductive rate of main Hawaiian Islands monk seals exceeded 0.70 for prime aged females (8–18 years). This is the highest reproductive rate reported for any of the Hawaiian monk seal breeding sites, illustrating the important role of the main Hawaiian Islands population in Hawaiian monk seal recovery.     

IDENTIFYING DIURNAL FORAGING HABITAT OF ENDANGERED HAWAIIAN MONK SEALS USING A SEAL-MOUNTED VIDEO CAMERA

The Hawaiian monk seal ( Monachus schauinslandi ) is thought to be a foraging generalist, preying on numerous species in diverse habitats of the subtropical Northwestern Hawaiian Islands. At the atoll of French Frigate Shoals, recent evidence of emaciation and low survival in monk seals prompted a search for their specific prey communities and foraging habitat.
A video camera (National Geographic Television's CRITTERCAM) fitted to 24 adult male seals documented benthic and demersal foraging on the deep slopes (50-80 m) of the atoll and neighboring banks. The number of bottom searches for prey was compared by year, time of day, type of bottom, individual seal, and length of bottom time. Analysis of variance identified a significant interaction of seal and bottom type, explaining 65% of the total variance. Seals fed on communities of cryptic fauna (fish and large invertebrates) in transitional "ecotone" regions of low relief where consolidated substrate, rubble, and talus bordered areas of sand. Independent areal surveys of bottom types throughout the atoll and neighboring banks suggest that the type of bottom selected as foraging habitat represents a relatively small percentage of the total benthic area available.     

HAWAIIAN MONK SEALS (MONACHUS SCHAUINSLANDI) FORAGING IN DEEP-WATER CORAL BEDS

Plans to harvest deep-water corals in the remote Northwestern Hawaiian Islands, close to populations of endangered Hawaiian monk seals ( Monachus schauinslandi ), have raised concerns about the seals' use of deep-water habitats. Movements and diving patterns of seals studied at French Frigate Shoals (FFS) Atoll indicated two areas where five males out of 33 instrumented seals dove deep enough (300–500 m) to encounter commercially sought deep-water corals. Submarine surveys conducted at each location found beds of gold ( Gerurdia sp.) and pink ( Corallium sp.) precious coral suggesting an overlap between the foraging habitat of some seals and the target of the coral fishery. Areas adjacent to the coral beds that were visually censused using submersibles showed significantly fewer precious corals. Precious coral beds were not found on previous submarine surveys at other regions around FFS, supporting the notion that seals were selecting the areas with corals as forage habitat. Five male seals were fitted with back-mounted video cameras to document feeding among precious corals. None of the five seals dove deep enough to encounter precious corals (>300 m). However, three of the seals visited beds of black coral ( Cirrhipdhes sp.) at shallower depths (∼ 80 m). One seal was observed revisiting the black coral beds on three successive nights to feed on fish hiding among the coral stems.     

Do monk seals exert top‐down pressure in subphotic ecosystems?

Patterns of subphotic fish assemblages on seamounts in the Northwestern Hawaiian Islands were identified and compared for potential structuring influences, including the bottom-up effects of regional oceanic productivity and top-down predation pressure exerted by visiting monk seals ( Monachus schauinslandi ). Patterns in fish size, density, and biomass were evaluated at the deep extreme (350–500 m) of the seals feeding range to avoid confounding effects of diverse shallow habitats ( e.g. , coral reefs). Fish number and size were used to calculate biomass density of the seamount fish assemblages that were then compared to the independent variables of summit depth, substrate type, relief, oceanic productivity, distance to seal colonies, and seal colony population. Only the variables of distance to seal colony and seal colony population were retained in a multiple regression model that explained 31% of the variance. Despite the presence of obvious regional differences in oceanic productivity, the overall patterns in the subphotic fish assemblages are better explained by the top-down hypothesis of predation pressure from monk seals.     

Biogeography and taxonomy of extinct and endangered monk seals illuminated by ancient DNA and skull morphology

Extinctions and declines of large marine vertebrates have major ecological impacts and are of critical concern in marine environments. The Caribbean monk seal, Monachus tropicalis, last definitively reported in 1952, was one of the few marine mammal species to become extinct in historical times. Despite its importance for understanding the evolutionary biogeography of southern phocids, the relationships of M. tropicalis to the two living species of critically endangered monk seals have not been resolved. In this study we present the first molecular data for M. tropicalis, derived from museum skins. Phylogenetic analysis of cytochrome b sequences indicates that M. tropicalis was more closely related to the Hawaiian rather than the Mediterranean monk seal. Divergence time estimation implicates the formation of the Panamanian Isthmus in the speciation of Caribbean and Hawaiian monk seals. Molecular, morphological and temporal divergence between the Mediterranean and “New World monk seals” (Hawaiian and Caribbean) is profound, equivalent to or greater than between sister genera of phocids. As a result, we classify the Caribbean and Hawaiian monk seals together in a newly erected genus, Neomonachus. The two genera of extant monk seals (Monachus and Neomonachus) represent old evolutionary lineages each represented by a single critically endangered species, both warranting continuing and concerted conservation attention and investment if they are to avoid the fate of their Caribbean relative.     

Movements and home ranges of monk seals in the main Hawaiian Islands

Hawaiian monk seals (Neomonachus schauinslandi) began recolonizing the main Hawaiian Islands (MHI) roughly 20 yr ago. The species’ abundance is still declining, but the subpopulation in the MHI is increasing by 6.5% per year. This difference may be due to differences in prey availability or habitat quality between the northwest (NWHI) and main Hawaiian Islands, which could be reflected in the movements and behavior of the seals. For example, foraging trip durations may be shorter in areas with higher forage quality. From 2007 to 2014 we deployed GPS phone tags on the islands of Molokai (n = 7), Kauai (n = 6), and Oahu (n = 6) to study movements and dive behavior. Foraging trips typically lasted 0.57 d (IQR: 0.34–0.83) and seals traveled 18 km (IQR: 10.2–30.5) per trip. Seals began benthic dives shortly after entering the water, with most dives to depths of 12–32 m. The median 95% and 50% kernel density isopleths for seals in the MHI were 149.2 km² and 23.2 km², respectively. The duration and distance of foraging trips in the MHI were shorter than that observed in other studies from the NWHI, suggesting that foraging habitat is currently better in the MHI.     

MORPHOMETRY, HEMATOLOGY, AND SERUM CHEMISTRY IN THE HAWAIIAN MONK SEAL (MONACHUS SCHAUINSLANDI)

We studied morphometric, hematology, and serum chemistry variables in 140 Hawaiian monk seals ( Monachus schauinslandi ) to establish normal baseline values for these variables among free-living seals. We compared seals at French Frigate Shoals (FFS), Midway Atoll (MID), and Pearl and Hermes Reef (PHR) because these subpopulations differ in their rates of population recovery. Dorsal standard length and axillary girth differed significantly between immature (1–4 yr old) and adult (≥5 yr old) seals among sex and island subgroups. Immature seals at FFS were shorter than those at MID and PHR; adult seals at FFS had smaller dorsal standard lengths and axillary girths compared to the other subpopulations. The differences in size were more pronounced among adult females. Significant differences were also found for hematology and serum chemistry variables among seals at FFS, MID, and PHR. Monk seals at FFS had an absolute lymphopenia and eosinopenia compared to those at MID and PHR, compatible with a stress response. Seals at FFS also had lower blood urea nitrogen than seals at PHR, and a lower plasma potassium than seals at MID or PHR. Monk seals had an absolute and relative eosinophilia compared to previously published values. Analysis of subpopulation differences is useful for population health assessment and for long-term monitoring of an endangered species.     

Seasonal testosterone pattern in Hawaiian monk seals (Monachus schauinslandi).

Blood samples from four captive male Hawaiian monk seals were collected at intervals of one month for one year for testosterone assay. Plasma testosterone concentrations, measured by radioimmunoassay, revealed a clear seasonal pattern. The lowest mean testosterone concentration (0.09 +/- 0.04 ng ml-1) occurred in January, and the highest (1.78 +/- 0.40 ng ml-1) in June. The seasonal occurrence of births and of injuries related to mating in wild populations of Hawaiian monk seals showed a distinct association with the period of high testosterone. This study supports other data that indicate that the Hawaiian monk seal is a seasonal breeder and is reproductively active for longer than monachine seals that live in higher latitudes.     

Ringed and bearded seal densities in the eastern Chukchi Sea, 1999–2000

Aerial surveys were conducted in 1999 and 2000 to estimate the densities of ringed (Phoca hispida) and bearded (Erignathus barbatus) seals in the eastern Chukchi Sea. Survey lines were focused mainly on the coastal zone within 37 km of the shoreline, with additional lines flown 148–185 km offshore to assess how densities of seals changed as a function of distance from shore. Satellite-linked time-depth recorders were attached to ringed seals in both years to evaluate the time spent basking on the ice surface. Haulout patterns indicated that ringed seals transitioned to basking behavior in late May and early June, and that the largest proportion of seals (60–68%) was hauled out between 0830 and 1530 local solar time. Ringed seals were relatively common in nearshore fast ice and pack ice, with lower densities in offshore pack ice. The average density of ringed seals was 1.91 seals km⁻² in 1999 (range 0.37–16.32) and 1.62 seals km⁻² in 2000 (range 0.42–19.4), with the highest densities of ringed seals found in coastal waters south of Kivalina and near Kotzebue Sound. The estimated abundance of ringed seals for the entire study area was similar in 1999 (252,488 seals, SE=47,204) and 2000 (208,857 seals, SE=25,502). Bearded seals were generally more common in offshore pack ice, with the exception of high bearded seal numbers observed near the shore south of Kivalina. Bearded seal densities were not adjusted for haulout behavior, and therefore, abundance was not estimated. Unadjusted average bearded seal density was 0.07 seals km⁻² in 1999 (range 0.011–0.393) and 0.14 seals km⁻² in 2000 (range 0.009–0.652). Levels of primary productivity, benthic biomass, and fast ice distribution may influence the distributions of ringed and bearded seals in the Chukchi Sea. Information on movement and haulout behavior of ringed and bearded seals would be very useful for designing future surveys.     

EFFECTS OF RESEARCH HANDLING ON THE ENDANGERED HAWAIIAN MONK SEAL

We examined the effects of research handling on free-ranging endangered Hawaiian monk seals, Monachus schauinslandi , by analyzing differences in subsequent year survival, migration, and condition between handled seals and controls during 1983–1998. Each of 549 handled seals was matched to a control seal of the same age, sex, location, and year. Handling included instrumentation with tel metry devices ( n = 93), blood sampling ( n = 19), and tagging ( n = 437). No significant differences were found between handled seals and their controls in one-year resighting rates, observed migration rates, or condition. Resighting rates of handled and control seals were high (80%-100%). Available sample sizes were sufficient to detect reasonably small (9%-20%) differences in resighting rates had they existed among instrumented or tagged seals and controls (α= 0.05, power = 0.90). Too few seals were captured for blood sampling to detect even large differences in their resighting rates. However, blood samples were drawn from most instrumented seals, and there was no indication that this larger group suffered harmful effects. Duration of restraint during flipper tagging had no effect on subsequent probability of resighting. Our analysis suggests that conservative selection procedures and careful handling techniques have no deleterious effects on Hawaiian monk seals.     

Modelled mid-trophic pelagic prey fields improve understanding of marine predator foraging behaviour

Biophysical interactions are influential in determining the scale of key ecological processes within marine ecosystems. For oceanic predators, this means foraging behaviour is influenced by processes shaping the distribution of prey. However, oceanic prey is difficult to observe and its abundance and distribution is regionally generalised. We use a spatiotemporally resolved simulation model to describe mid-trophic prey distribution within the Southern Ocean and demonstrate insights that this modelled prey field provides into the foraging behaviour of a widely distributed marine predator, the southern elephant seal. From a five-year simulation of prey biomass, we computed climatologies of mean prey biomass (average prey conditions) and prey biomass variability (meso-scale variability). We also compiled spatially gridded metrics of seal density and diving behaviour from 13 yr of tracking data. We statistically modelled these metrics as non-linear functions of prey biomass (both mean and variability) and used these to predict seal distribution and behaviour. Our predictions were consistent with observations (R²_adj = 0.23), indicating that seals aggregate in regions of high mesoscale activity where eddies concentrate prey. Here, seals dived deeper (R²_marg = 0.12, R²_cond = 0.51) and spent less time hunting (R²_marg = 0.05, R²_cond = 0.56), likely targeting deep but profitable prey patches. Seals generally avoided areas of low eddy activity where prey was likely dispersed. Most seals foraged south of the Subantarctic Front, despite north of the front exhibiting consistently high simulated prey biomasses. This likely reflects seal prey or habitat preferences, but also emphasises the importance of mesoscale prey biomass variability relative to regionally high mean biomass. This work demonstrates the value of coupling mechanistic representations of prey biomass with predator observations to provide insight into how biophysical processes combine to shape species distributions. This will be increasingly important for the robust prediction of species’ responses to rapid system change.     

CONCENTRATIONS OF ESTRONE SULFATE and PROGESTERONE IN PLASMA and SALIVA, VAGINAL CYTOLOGY, and BIOELECTRIC IMPEDANCE DURING THE ESTROUS CYCLE OF THE HAWAIIAN MONK SEAL (MONACHUS SCHAUNSLAnDI)

Blood and saliva samples, vaginal swabs, and bioelectric impedance measurements were collected 1–3 times per week from a captive adult female Hawaiian monk seal ( Monacbus schauinslandi ) during the spring and summer of 1991. During 1992 saliva only was collected on average 4 times weekly. Concentrations of progesterone and estrone sulfate, and the periodic appearance of cornified epithelial cells from the vagina indicated consecutive estrous cycles ranging 35 ± 3 days in duration. Progesterone concentrations in plasma and saliva had a correlation of 0.868, while estrone sulfate had a correlation of 0.982. Bioimpedance of the whole seal body resulted in a pattern similar to the estrone sulfate concentrations, but shifted forward by 2 d. Bioimpedance of the upper vaginal tissues paralleled the estrone sulfate pattern while the bioimpedance of the lower reproductive tract fluctuated without a distinct pattern. The luteal phase ranged 17–20 d and the follicular phase was 15–18 d in length. These results indicate that: (1) the Hawaiian monk seal, in captivity, is a polyestrous seal, in contrast to other phocid seals that have been studied; (2) salivary concentrations of estrogen and progesterone may provide an accurate, less-invasive method of monitoring reproductive hormones in captive Hawaiian monk seals; and (3) vaginal cytology and bioelectric impedance reflect physiological changes associated with the estrous cycle.     

Extremely low genetic diversity in the endangered Hawaiian monk seal (Monachus schauinslandi)

Hunted to near extinction in the late 19th century, the endangered and endemic Hawaiian monk seal (Monachus schauinslandi) exhibits low variation at all molecular markers tested to date. Here we confirm extreme paucity of genetic diversity, finding polymorphisms at only 8 of 154 microsatellite loci tested (143 novel species-specific loci, 10 loci from Antarctic seals, and 1 previously characterized locus). This screening revealed unprecedentedly low levels of allelic diversity and heterozygosity (A = 1.1, H(e) = 0.026). Subsequent analyses of 2409 Hawaiian monk seals at the 8 polymorphic loci provide evidence for a bottleneck (P = 0.002), but simulations indicate low genetic diversity (H(e) < 0.09) prior to recorded human influence. There is little indication of contemporary inbreeding (F(IS) = 0.018) or population structure (K = 1 population). Minimal genetic variation did not prevent partial recovery by the late 1950s and may not be driving the current population decline to approximately 1200 seals. Nonetheless, genotyping nearly every individual living during the past 25 years sets a new benchmark for low genetic diversity in an endangered species.     

Nutrient, fatty acid, amino acid and mineral analysis of natural prey of the Hawaiian monk seal, Monachus schauinslandi.

Proximate nutrients, gross energy content, mineral, amino acid and fatty acid composition were determined for teleost, cephalopod and crustacean prey of the Hawaiian monk seal. Crude protein was highest in the octopus, Octopus cyanea (80.0%), crude fat was highest in the Muraenid teleost, Gymnothorax eurostus (14.1%), whereas crude ash was highest in the lobster, Panulirus marginatus (11.6%). Gross energies ranged from 4.0 +/- 0.01 kcal g-1 in the Labrid teleost Bodianus bilulunatus to 6.0 +/- 0.12 kcal g-1 in the moray eel, Gymnothorax undulatus. Essential amino acids occurred in lower concentrations as a percentage of the total amino acids (35.8 +/- 2.6%) than non-essential amino acids (64.2 +/- 2.6%), but the ratio of individual amino acids to total amino acid concentrations were similar to those required by some monogastric terrestrial species and fingerling salmon. The fatty acid concentrations varied widely among species (range = 1.2-16.5 mg 100 mg-1); however, the teleosts had higher total fatty acids than the non-teleosts. This study indicates that, from a nutritional standpoint, some prey may be more beneficial to the Hawaiian monk seal; however, these prey are not necessarily the most abundant or available to some populations of the monk seal.     

ABUNDANCE OF RINGED SEALS (PUSA HISPIDA) IN THE FJORDS OF SPITSBERGEN, SVALBARD, DURING THE PEAK MOLTING PERIOD

Ringed seal (Pusa hispida) abundance in Spitsbergen, Svalbard, was estimated during the peak molting period via aerial, digital photographic surveys. A total of 9,145 images, covering 41.7%–100% of the total fast‐ice cover (1,496 km²) of 18 different fjords and bays, were inspected for the presence of ringed seals. A total of 1,708 seals were counted, and when accounting for ice areas that were not covered by images, a total of 3,254 (95% CI: 3,071–3,449) ringed seals were estimated to be hauled out during the surveys. Extensive behavioral data from radio‐tagged ringed seals (collected in a companion study) from one of the highest density fjords during the molting period were used to create a model that predicts the proportion of seals hauled out on any given date, time of day, and under various meteorological conditions. Applying this model to the count data from each fjord, we estimated that a total of 7,585 (95% CI: 6,332–9,085) ringed seals were present in the surveyed area during the peak molting period. Data on interannual variability in ringed seal abundance suggested higher numbers of seals in Van Keulenfjorden in 2002 compared to 2003, while other fjords with very stable ice cover showed no statistical differences. Poor ice conditions in general in 2002 probably resulted in seals from a wide area coming to Van Keulenfjorden (a large fjord with stable ice in 2002). The total estimated number of ringed seals present in the study area at the time of the survey must be regarded as a population index, or at least a minimum estimate for the area, because it does not account for individuals leaving and arriving, which might account for a considerable number of animals. The same situation is likely the case for many other studies reporting aerial census data for ringed seals. To achieve accurate estimates of population sizes from aerial surveys, more extensive knowledge of ringed seal behavior will be required.     

Dramatic shifts in Hawaiian monk seal distribution predicted from divergent regional trends

Total estimated abundance of Hawaiian monk seals was just 1,161 individuals in 2008 and this number is decreasing. Most monk seals reside in the remote Northwestern Hawaiian Islands (NWHI) where the decline is approximately 4%/yr, whereas relatively fewer seals currently occupy the main Hawaiian Islands (MHI). It is widely accepted that the MHI population is increasing, although there are no formal estimates of total abundance, population growth rate or vital rates. This lack of information has hampered efforts to anticipate future scenarios and plan conservation measures. We present the first estimates of MHI monk seal survival and age‐specific reproductive rates. Using these rates, a conservative estimate of current MHI abundance and a previously published stochastic simulation model, we estimate the MHI population growth rate and projected abundance trend. Analogous estimates for the NWHI are derived from a much richer data set. Estimated survival from weaning to age 1 yr is 77% in the MHI, much higher than recent NWHI estimates ranging from 42% to 57%. Moreover, MHI females begin reproducing at a younger age and attain higher birth rates than observed in the NWHI. The estimated MHI intrinsic rate of population growth is 1.07 compared to a 0.89–0.96 range in the NWHI. Assuming an initial abundance of 152 animals in the MHI, projections indicate that if current demographic trends continue, abundance in the NWHI and MHI will equalize in approximately 15 yr. These results underscore the imperative to mitigate the NWHI decline while devoting conservation efforts to foster population growth in the MHI, where documented threats including fishery interactions, direct killing, and disease could rapidly undo the current fragile positive trend.     

Temporal variation in distribution and density of ice-obligated seals in western Hudson Bay, Canada

Recent unidirectional climatic trends and changes in top predator population ecology suggest that long-term modifications may be happening in Hudson Bay, Canada. Effects of such changes on ice-obligated seal populations are expected but long-term studies are required to differentiate climate-induced changes from natural variation. We conducted strip-transect surveys in late spring in 1995–1997, 1999–2000 and 2007–2008 to estimate distribution, density and abundance of ice-obligated ringed (Phoca hispida) and bearded (Erignathus barbatus) seals in western Hudson Bay. When hauled out, ringed seals preferred land-fast and consolidated pack ice, whereas bearded seals preferred unconsolidated pack ice. Bearded and ringed seal density estimates varied from 0.0036 to 0.0229 seals/km² of ice and from 0.46 to 1.60 seals/km² of ice, respectively. Strong inter-annual variations were recorded in the abundance estimates of both species, with the largest abundance estimates in 1995 (104,162 and 1,494 ringed and bearded seals, respectively) and the lowest in 2008 for ringed seals (33,701) and 1997 for bearded seals (278). A sine function best described seal density estimates in western Hudson Bay and suggested a decadal cycle. Previous studies that reported low ringed seal demographic parameters in the 1990s and a recovery in the 2000s supported our interpretation of the survey results. We discuss our results in the context of climate warming and suggest that a long-term decline in ice-obligated seal density estimates may overlay a possible natural decadal cycle.     

Distribution and abundance of spotted seals Phoca largha and ribbon seals Phoca fasciata in the southern Sea of Okhotsk

The distribution and abundance of spotted seals (Phoca largha) and ribbon seals (Phoca fasciata) were assessed in March and April, 2000, by aerial line-transect surveys along the southern edge of the pack ice off the coast of Hokkaido (southern Sea of Okhotsk), Japan. Five hundred and seventeen spotted seals and 107 ribbon seals were found on the total 2944 km survey line. Total abundance was estimated to be 13 653 spotted (95% CI = 6167–30 252) and 2260 ribbon seals (95% CI = 783–6607) in March, and 6545 spotted (95% CI = 3284–815 644) and 3134 ribbon seals (95% CI = 1247–17 802 512) in April. The pack ice area off Hokkaido had higher densities (0.54 seals km^–2 and 0.58 seals km^–2 in March and April, respectively) of spotted seals than those reported in eastern Sakhalin, whereas densities (0.09 seals km^–2 in March and 0.28 seals km^–2 in April) of ribbon seals were lower than those in eastern Sakhalin. The large number of spotted seal pups suggests that the study area is an important breeding center. A greater number of female spotted seals with pups tended to be found in the center of larger and rougher floes than in other categories, and they were more abundant in stable pack ice areas. Observations of ribbon seals were limited because the survey period preceded the peak of pupping season. Ribbon seal surveys were also hampered by the inability to fly over the main breeding area between the Shiretoko Peninsula and Kunashiri Island.     

USING TREND INDICES FOR ENDANGERED SPECIES

Initial appraisals of the status of endangered large-mammal populations may have to depend on indices of population trend. Such indices may possibly be improved by using auxiliary variables. Various models were studied for populations of the Florida manatee (Trichechus manatus latirostris), Yellowstone grizzly bear (Ursus arctos horribilis), and Hawaiian monk seal (Monachus schauinslandi). Several criteria for checking validity of the fitted models were considered, and the simple R² criterion appears to provide useful comparisons. Multiple regression models overestimated the rate of change of the East Coast manatee population as determined from three other sources (a covariance model, a non-linear model, and the rate estimated from reproductive and survival data). A multiple regression model for grizzly bears using three auxiliary variables exhibited a fairly high R² (0.84) and appeared to provide a better fit than did a non-linear model. A beach count index for Hawaiian monk seals seemed to be unreliable for year-to-year comparisons in contrast to total population counts and estimates from a capture-recapture method. The use of auxiliary variables for checking and improving trend index data appears feasible and well worthwhile.     

SURVIVAL RATES FOR THE HAWAIIAN MONK SEAL (MONACHUS SCHAUINSLANDI)

Abstract: Endangered Hawaiian monk seal ( Monachs schauinslandi ) pups at all the major breeding islands in the Northwestern Hawaiian Islands have been tagged since the early 1980s. Pups were double flipper tagged as soon as possible post-weaning. With few exceptions, an extensive tag resighting effort was conducted annually at the same islands. These resighting data were used to estimate seal survival rates from the time of tagging to age one at all locations using the ratio of seals alive in the second year to number of pups tagged. These survival rates among the islands, from weaning to age one, averaged over the years of the study, ranged from 0.80 to 0.90. For young seals over age one, capture-recapture methods were used to calculate survival pooled through several years, and these rates ranged from 0.85 to 0.98. At French Frigate Shoals and Laysan Island, the higher numbers of tagged pups allowed separate estimates of male and female survival to be calculated. These rates suggested that survival of immature females was better than males. Beginning in 1989, survival of immature seals at French Frigate Shoals declined sharply.