IMPROVED BODY CONDITION OF WEANED HAWAIIAN MONK SEAL PUPS ASSOCIATED WITH EL Niño EVENTS: POTENTIAL BENEFITS TO AN ENDANGERED SPECIES

Efforts to enhance recovery of endangered Hawaiian monk seals ( Monachus schauinslandi ) require an understanding of factors influencing population dynamics. This study examines relationships between body condition and survival of monk seal pups at French Frigate Shoals and Laysan Island and El Niño events. Girth measurements and mass estimates were used as indicators of pup body condition, and survival was evaluated from weaning to age 2. Linear models and logistic regression analysis were used to evaluate El Niño effects. Temporal trends in mean girth, mass, and survival were identified at both study sites. After accounting for temporal trends, girths were 3.7 cm and 2.7 cm greater during El Niño years at French Frigate Shoals and Laysan Island, respectively. Average mass estimates were significantly greater during El Niño years at French Frigate Shoals (2.6 kg increase), but were not significantly different at Laysan Island (1.8 kg increase). Weaned pups born at French Frigate Shoals during El Niño years survived significantly better, but this effect was not detected at Laysan Island. El Niño events probably affect pup condition and survival, but these parameters need to be monitored during future El Niño events to clarify these relationships.     

BODY SIZE,SURVIVAL, AND DECLINE OF JUVENILE HAWAIIAN MONK SEALS,MONACHUS SCHAUINSLANDI

In the mid 1980s half of the entire Hawaiian monk seal species was located at French Frigate Shoals, and this colony may have reached environmental carrying capacity. Since 1989 this colony has declined by 55%, primarily from poor juvenile survival. Only 8%–25% of weaned pups have survived to age 2 during this period, compared to at least 80% between 1984 and 1987. We characterize (1) this drop in survival, (2) a strong, but variable, correlation between survival and size at weaning, (3) interisland differences in size from weaning to age 2 in 1991 through 1993, and (4) decadal-scale changes in mean measures of size at weaning. We compare observations at French Frigate Shoals with the colony at Laysan Island where abundance is well below historical levels and was therefore expected to be below carrying capacity. At Laysan Island juvenile survival has also been poor (30%-70%), and the size of weaned pups has decreased during the past decade. Our hypothesis is that poor survival may be due to reduced prey availability at both sites, but particularly at French Frigate Shoals. Evidence to support this hypothesis includes a large increase in total abundance at French Frigate Shoals; reduced size of weaned pups; decreased survival, emaciation, and slower growth rates of juvenile seals; and declining primary productivity. Regardless of cause, the immediate consequences at French Frigate Shoals and Laysan Island, and for the species overall, will likely be poor recruitment and productivity. Because the decline is still in progress, the ultimate consequences for the species' viability are of great concern in light of its already low abundance.     

Distribution and frequencies of shark-inflicted injuries to the endangered Hawaiian monk seal (Monachus schauinslandi)

I analyzed the frequencies of wounds inflicted by sharks to the endangered Hawaiian monk seal Monachus schauinslandi at three colonies in the Northwestern Hawaiian Islands between 1990 and 2000. I applied specific criteria typical of shark bites to all injuries to avoid bias. Pooling data from all years and the three colonies revealed several patterns. Sharks injured more pups (nursing and weaned) and juveniles (1–2 years old) than adults and subadults. More female pups and male juveniles than any other size classes were injured by sharks. Almost all (97%) of the injured pups were from French Frigate Shoals. More juveniles than expected were wounded at Laysan Island and Lisianski Island. Most shark wounds were between the diaphragm and the pelvic girdle, but pups were bitten most often just behind the pelvis compared with adults who were bitten more often near the head and neck.     

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.     

Estimating the carrying capacity of French Frigate Shoals for the endangered Hawaiian monk seal using Ecopath with Ecosim

The carrying capacity of the French Frigate Shoals (FFS) region for the endangered Hawaiian monk seal was appraised using an updated version of the original FFS Ecopath model ( Polovina 1984 ). Model parameters were updated using recent literature, and data from surveys of the seal population and its bottom‐associated prey. Together they produced a static mass balance model for 1998 when the prey surveys began. The Ecopath‐estimated monk seal biomass was 0.0045 t/km², which was in close agreement with the biomass calculated from monk seal field beach counts (0.0046 t/km²). Model simulations through time were done in Ecosim using the Ecopath balanced model and included fisheries data time series from 1998 to 2008. Monk seal biomass declined concurrently with decreases in benthic bottomfish biomass, which were influenced by large‐scale changes in the environment of the North Pacific. This model scenario was extended from 2010, when the last permitted fishery in the Northwestern Hawaiian Islands was closed, through to 2040, assuming a constant environmental signal. Model results for this time period did not show a recovery of monk seals that exceeded the initial 1998 model biomass levels, highlighting the importance of including environmental variability in estimates of monk seals recovery at FFS.     

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.     

A COPROLOGICAL SURVEY OF HELMINTH PARASITES OF THE HAWAIIAN MONK SEAL FROM THE NORTHWESTERN HAWAIIAN ISLANDS

In conjunction with an Hawaiian monk seal, Monachus schauinslandi , breeding enhancement program, fecal samples were examined for the presence of helminth parasite products (eggs, larvae, and proglottids). Eighty-two samples were examined from four island groups in the northwestern Hawaiian Islands: 38 from Kure Atoll, 32 from French Frigate Shoals, 11 from Laysan Island, and one from Lisianski Island. Identified helminth products include those of Diphyllobothrium sp., D. cameroni, Contracaecum turgidum, Anisakis sp., Corynosoma rauschi , and Trematoda. In addition, eggs from Halarachne laysanae (Atari) were found in 3 samples. Eggs of C. turgidum were present in the highest percentage (>64%) of samples from all locations. Eggs of Anisakis sp. and Trematoda are reported from this host for the first time.     

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.     

First-year survival of southern elephant seals,Mirounga leonina,at sub-Antarctic Macquarie Island

Juvenile seals branded on the isthmus of Macquarie Island as pups displayed a high degree of philopatry. They returned more often and in greater densities to the northern third of the island within 10 km of their birth sites. Juvenile seals were observed to haul out more frequently and in greater numbers on the east coast as opposed to the west. Juvenile seals typically hauled out on two occasions, once during the winter, and once to moult. The probability of recapturing (resighting) branded and tagged seals was greater during the mid-year haulout. First-year survival estimates were obtained from searches of all Macquarie Island beaches for marked (branded and tagged) seals. From a branded population of 2000 seals, 897 were known to be alive at age 1 year, and minimum first-year survival was calculated at 44.85%. To this minimum estimate was added the number of seals overlooked during systematic and standardised searches of the island, and a revised estimate of 65.60% was calculated. Survival rates calculated using a custom model and a conventional mark-recapture model (MARK) were compared and no differences detected. Actual survival data and probability of sighting estimates were included in the revised estimate of first-year survival of southern elephant seals at Macquarie Island. There were no differences in the number of surviving males and females. Accepted: 25 October 1998     

Temporal and spatial variation in age-specific survival rates of a long-lived mammal, the Hawaiian monk seal

Estimates of variability in pinniped survival rates are generally based on observations at single sites, so it is not certain whether observed rates represent the whole population. Here, we provide a comprehensive analysis of spatio-temporal variation in age-specific survival rates for endangered Hawaiian monk seals (Monachus schauinslandi) based on capture-recapture analyses of more than 85% of the pups weaned in this population over the last two decades. Uniquely, these data have been collected from six subpopulations, encompassing all major breeding sites across its 1800 km long core range. Analyses of individual subpopulations revealed similar patterns in age-specific survival, characterized by the relatively low survival rates from weaning to 2 years of age, intermediate rates to 4 years of age, and then by relatively high 'mature' survival rates until 17 years of age, after which a senescent decline was observed. Juvenile, subadult and adult survival rates all varied significantly over time. Trends in survival among subpopulations were coherent with their relative geographical positions, suggesting regional structuring and connectedness within the archipelago. Survival rates for different age classes tended to be positively correlated, suggesting that similar factors may influence the survival for seals of all ages.     

Colonization of the Hawaiian Archipelago via Johnston Atoll: a characterization of oceanographic transport corridors for pelagic larvae using computer simulation

Larval transport between Johnston Atoll and the Hawaiian Archipelago was examined using computer simulation and high-resolution ocean current data. The effects of pelagic larval duration and spawning seasonality on long-distance transport and local retention were examined using a Lagrangian, individual-based approach. Retention around Johnston Atoll appeared to be low, and there appeared to be seasonal effects on both retention and dispersal. Potential larval transport corridors between Johnston Atoll and the Hawaiian Archipelago were charted. One corridor connects Johnston Atoll with the middle portion of the Hawaiian Archipelago in the vicinity of French Frigate Shoals. Another corridor connects Johnston Atoll with the lower inhabited islands in the vicinity of Kauai. Transport appears to be related to the subtropical countercurrent and the Hawaiian Lee countercurrent, both located to the west of the archipelago and flowing to the east. A new analytical tool, termed CONREC–IRC is presented for the quantification of spatial patterns.     

Sexual reproduction in Hawaiian Acropora

Sexual reproductive maturity was evaluated in Acropora valida, Acropora cytherea and Acropora humilis at French Frigate Shoals, Northwest Hawaiian Islands during two consecutive summers. Acropora valida gonads matured at different rates in different habitats in both years. Spawning was inferred by the sudden disappearance of gonads of mature size. Histological sections of fertile polyps confirmed the maturity of gonads prior to spawning. An isolated colony of A. humilis spawned in early summer. Strong indications of sexually mature colonics of A. cytherea exist, but clear temporal patterns were not apparent. Lunar period of spawning in reef flat A. valida and A. humilis differed from that reported for these species from other regions. The three species did not overlap in time of spawning. Previous ideas concerning the allochthonous origin of larval recruits, as well as the absence of Acropora from high islands of the Hawaiian chain, are re-evaluated in light of new evidence for sexual reproductive capacity by native populations.     

FORAGING OF JUVENILE MONK SEALS AT FRENCH FRIGATE SHOALS, HAWAII

Emaciation and poor survivorship of juvenile Hawaiian monk seals at French Frigate Shoals atoll prompted a study of their foraging, using video camera technology ( crittercam ). Nine juveniles between the ages of 1 and 3 yr (six males, three females) were fitted with crittercam to identify their foraging habitat and feeding behavior. All feeding was directed at small (≤ 10 cm), cryptic, benthic prey. Older seals (ages 2 and 3), varied in their foraging intensity with most of their attention directed at shallow atoll depths (10–30 m). In contrast, the three yearlings focused all their feeding in the sand fields (50–100 m) on the atoll's outer slope. Bottom trawls were used to assess the prey abundance of the sand habitat and found 70% of the numerical catch was flounder ( Bothidae ). Extrapolating the yearlings' prey capture rate (0.13/min, derived from the crittercam video) over their total bottom time yielded an estimated 1–1.3 kg/day of flounder. The mean size of flounder (5 ± 1.7 cm) caught in the bottom trawls was close to the size at which larval flounder settle from the plankton (3 cm), suggesting that localized changes in oceanography could directly impact the seals' prey supply. Extensive use of sand communities by young seals may be the strongest link yet identified between juvenile survivorship and oceanographic dynamics.     

REPRODUCTIVE PATTERNS OF THE HAWAIIAN MONK SEAL

We evaluated reproductive patterns of the Hawaiian monk seal ( Monachus schauinslandi ) using a combination of fitted age-specific reproductive curves and analysis of reproductive patterns of individual females. We review the difficulties inherent in the acquisition and modeling of reproductive data with emphasis on the significance of reproductive senescence to populations with dissimilar age/sex compositions. Validation of the fitted reproductive parameters was accomplished by Monte Carlo sampling of parameter distributions to compare the expected number of pups with the observed production. Although the fitted reproductive functions appear to provide an acceptable fit to the raw reproductive data, we found that the fitted curves did a poor job of predicting the actual pup production in individual years because of high variability among years. To further verify, and elaborate on, the patterns in the pooled (multi-seal, and multi-year) rates, we examined attributes of the reproductive performance of individual seals. The attributes included age of primiparity, reproductive rates computed over several age ranges, and the relationship between reproductive performance and seal longevity. Analysis of individual seal patterns reinforced the conclusion that reproductive senescence is operative in monk seal populations.     

Unmarked individuals in mark–recapture studies: Comparisons of marked and unmarked southern elephant seals at Marion Island

The presence of unmarked individuals is common in mark–recapture study populations; however, their origin and significance in terms of population dynamics remain poorly understood. At Marion Island, southern Indian Ocean, where virtually all southern elephant seal Mirounga leonina pups born annually (1983–2008) were marked in a long‐term mark–resight study, large numbers of unmarked seals occur. Unmarked seals originate either from marker (tag) loss or from immigration. We aimed to identify patterns in the occurrence of marked and unmarked individuals that will allude to the possible origin and significance of the untagged component of the population, predicting that tag loss will add untagged seals to mainly adult age categories whereas migrating untagged individuals will be mostly juveniles. We fitted a generalized linear model using the factors month, year and age‐class to explain the relative abundance of untagged seals (tag ratio) from 1997 to 2009. Site usage of untagged seals relative to tagged seals was assessed using a binomial test. Untagged seals, predominantly juveniles, were present in the highest proportions relative to tagged seals during the winter haulout (tagged seals/total seals less than 0.3) and the lowest proportion (approximately 0.5) during the female breeding haulout, increasing in relative abundance from 1997 to 2009. Untagged seals were distributed evenly across suitable haulout sites while tagged seals displayed high local site fidelity and occurred in greater numbers at or near large breeding beaches. Untagged seals are considered to be mostly migrant seals that disperse from other islands within the southern Indian Ocean and haul out at Marion Island during non‐breeding haulouts in particular. Some of these seals immigrate to the breeding population, which can be a key component of the local population dynamics. We emphasize the need for mark–recapture studies to evaluate the role of the unmarked component of a population, thereby inducing a more confident estimation of demographic parameters from the marked sample.     

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.     

168 Cell Wall Proteomics of the Green Alga Haematococcus Pluvialis (Chlorophyceae)

Coral reefs of US-held islands in the central Pacific Ocean are among the most pristine in the world and represent over 93% of the reef systems under United States jurisdiction. The remote location of many islands has limited past algal research, resulting in incomplete understanding of species diversity, quantity, and ecology. Starting in 2000, the Coral Reef Ecosystem Investigation (CREI) began rapid ecological assessments on many Pacific island reefs to monitor ecological changes in reef biota over time. During the past year, algal efforts have concentrated on the French Frigate Shoals (Northwestern Hawaiian Islands) where we have increased the number of algal species reported by 1000%. Additionally, species new to science, including Acrosymphyton brainardii and Scinaia huismanii, have been described. Quantitative field sampling using a photoquadrat method is revealing species of the green algae Halimeda and Microdicyton to be ecological dominants in many areas during late summer/early autumn. Preliminary analyses with Primer software show species composition and abundance of all benthic organisms to differ significantly between most field sites sampled. Additional benthic habitat mapping of Pacific island reefs by CREI researchers is breaking the long-held paradigm that macroalgal cover is minimal in healthy tropical reef systems. Videotape analyses of benthic communities often find over 50% algal cover from 1 to 20 meter depths in many locations. Common ratios of macroalgae, turf algae, and crustose coralline algae to corals, other benthic organisms and substrate types on US Pacific reefs are being calculated for the first time.     

Morphometrics,body condition,and growth of the ringed seal (Pusa hispida saimensis) in Lake Saimaa: Implications for conservation

Morphometrics and growth of the critically endangered Saimaa ringed seal (Pusa hispida saimensis), which inhabits a freshwater lake in Finland were studied using data from 344 carcasses. This study presents the first detailed information on ringed seal pup growth and body condition from birth to the age of one year. Newborn pups were on average 68 cm long and weighed 5 kg. Pups attained the majority of their first year growth during the nursing period. Body condition and growth declined after weaning, but pups recovered from postweaning nutritional deprivation at the age of 8 mo. The seals achieved their maximum body length, girth, and mandible size around the age of 4 yr, and asymptotic body mass two years later. Baculum growth indicated that males reached sexual maturity at age 5–6 yr. The Saimaa ringed seals' asymptotic body length and mass were 132 cm and 59 kg, respectively, which is similar to medium sized marine ringed seals. Saimaa ringed seals' survival to adulthood is extremely low due to high bycatch mortality; furthermore climate change may hamper pup growth and elevate mortality. Therefore we recommend continuation of fishing closures to improve pup survival.     

BODY TEMPERATURE IN THE NEWBORN SOUTHERN ELEPHANT SEAL, MIROUNGA LEONINA, AT MACQUARIE ISLAND

Newborn southern elephant seal pups were reported by Laws (1953) to be "to some extent poikilothermic at birth." Rectal temperatures of known age southern elephant seal pups were recorded during the 1985 pupping season at Macquarie Island. The mean pup rectal temperature was found to be 381°C ± 0.1°C SEM ( n = 131, range = 36.5°-39.1°C). Pups at two hours, six hours, and one day after birth had significantly higher rectal temperatures than pups two, three, or four days of age. Rectal temperatures of neonatal southern elephant seals were within the range observed for other pinnipeds, (but never as low as the 31°C previously observed for southern elephant seals at Signy Island in 1953). A significant though weak positive correlation was found between pup temperature and body weight. However, no correlation between pup temperature and age or any environmental factor was found. These observations demonstrated that southern elephant seal pups at Macquarie Island are homeothermic, rather than heterothermic from birth.