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.     

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.     

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.     

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.     

ANALYSIS OF CORPORA LUTEA TO ESTIMATE REPRODUCTIVE CYCLES OF WILD HAWAIIAN MONK SEALS (MONACHUS SCHAUINSLANDI)

Ovaries from 14 female Hawaiian monk seals, O-24 yr of age, collected from the northwestern Hawaiian Islands were analyzed by gross and histological examination for the presence of corpora hemorrhagica (CH), lutea (CL), albicantia (CA), and dominant follicles. Seven seals, 5-24 yr of age, were mature females, and another seven seals, O-5 yr of age, were immature based on the presence or absence of CL and CA in the ovaries. Two 5-yr-old seals did not have CL and CA, but one J-yr-old and both 6-yr-old seals had CL or CA. In addition, the weight of the more active ovary increased at 5 yr of age, indicating that puberty in these female Hawaiian monk seals occurred about 5 yr of age. The total number of CL and CA was significantly correlated to the ovarian weight, but not age of the seals, indicating that visible CA of Hawaiian monk seals probably degenerate within a year. If an ovulation resulted in pregnancy, we assumed that the resulting CL from that pregnancy persisted throughout gestation and regressed approximately 2 yr after ovulation. However, the CL of pregnancy were not easily differentiated from cyclic CL that did not result in pregnancy. Four of seven mature seals were clearly polyestrous from the analyses of their ovaries and reproductive histories. The remaining three seals were pregnant in previous consecutive years, and we could not positively determine whether they were also polyestrous. The ovaries from Hawaiian monk seals had unusually large, rounded granular cells containing the aging pigment, lipofuscin. These cells were located between loose fibrous tissues in all regressing CL and CA but were not present in any CH.     

ANNUAL REPRODUCTIVE CYCLE OF THE FEMALE HAWAIIAN MONK SEAL (MONACHUS SCHAUINSLANDI)

Abstract: The annual reproductive cycle is described for the adult female Hawaiian monk seal ( Monachus schauinslandi ) from data collected at Laysan Island (1982–1991) and Lisianski Island (1982–1983) in the Northwestern Hawaiian Islands. Pupping, lactation, weaning, and molting were directly observed, while mating was rarely observed and was, therefore, inferred from the occurrence of mounting injuries and from adult male and female association patterns. Pooled birth rates during the study period were 0.544 for all adult-sized females and 0.675 for females parous in earlier years. For parturient females, pupping peaked in late March and early April, weaning in May, mounting injuries in May and June, and molting in July. For non-parturient females, the median mounting injury and molting dates occurred 17 and 28 days earlier, respectively. Pupping date set the timing of subsequent events in the annual cycle, but the timing of those events was adjusted by loss of the pup or poor physical condition of the female. Individual pupping patterns varied widely. The mean interval for births in consecutive years was 381 days; females that pupped in consecutive years gave birth later each season. Conversely, females who skipped a year or more gave birth earlier their next pupping season.     

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.     

USE OF DISCOVERY CURVES TO ASSESS ABUNDANCE OF HAWAIIAN MONK SEALS

We investigated the pattern of first sighting of individual seals over the course of a field season, or the "discovery curve," as a means for estimating abundance of the endangered Hawaiian monk seal, Monachus schauinslandi . We empirically derived a criterion to determine whether or not total enumeration had been accomplished at a given site and year. When greater than 100-h field effort was expended without a new individual being identified, we concluded that total enumeration was likely achieved. To evaluate the potential for estimating abundance through extrapolation of nonlinear asymptotic functions fitted to discovery curves, we conducted simulations under a range of capture probability scenarios, including some based on observed individual variability in monk seal sighting frequencies. We demonstrated that if capture heterogeneity existed among individuals, the fitted asymptotes tended to yield biased estimates of abundance. Moreover, the levels of bias and uncertainty tended to increase inversely with the proportion of the population identified. While extrapolation shows little promise for generating unbiased abundance estimates, discovery curves have practical appeal for determining whether total enumeration has been achieved, and for optimizing field effort allocation. This is especially true for relatively small, closed populations of marked individuals.     

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.     

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.     

CHANGES IN MORPHOLOGY WITH AGE IN MEDITERRANEAN MONK SEALS (MONACHUS MONACHUS)

In order to describe the pelage and external appearance of the Mediterranean monk seal ( Monachus monachus ) and prepare an age-sex classification guide, 120 seals from the Cabo Blanco Peninsula colony in the Western Sahara/Mauritania were periodically photographed between 1993 and 1996. Analysis of the pigmentation pattern, pelage color and pattern of natural markings of each seal established 48 phenetic types, which were, in turn, arranged in five groups (morphological classes) with multivariate cluster analysis (UPGMA). The variables that best define these groups are size and color. The variation related to relative size, sex and age of 26 identified seals monitored over three years showed that: (1) after each molt, external appearance varied considerably only in non-adults, while adult appearance was invariable; (2) marked sexual dimorphism exists in adults' external appearance; and (3) a remarkable similarity of adult and neonate pelage exists. With these results, we propose an age-sex classification guide to facilitate monk seal identification in the field.     

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.     

PUP SURVIVAL IN THE MEDITERRANEAN MONK SEAL (MONACHUS MONACHUS) COLONY AT CABO BLANCO PENINSULA (WESTERN SAHARA-MAURITANIA)

We surveyed pup survival in Mediterranean monk seals ( Monachus monachus ), at Cabo Blanco Peninsula (Western Sahara-Mauritania) colony from May 1993 to December 1997. This species breeds and hauls out on beaches inside two main caves. During the study period we detected a total of 93 pups that died or disappeared. The survival rate of 84 pups through the age of first moult (approximately two months) was 0.47. This value is similar to those reported for other pinnipeds breeding in caves but lower than for those breeding on open beaches. Mortality varied seasonally and appeared to increase as a result of storms, large ocean swells, and high tides. Mother-pup pair separation (and resulting pup starvation) and physical injury caused by impact against the rock walls of the cave and cliffs were established as the causes of most deaths. Beach surface area inside the caves also appeared to be a mediating factor in the effects of sea conditions. High pup mortality may be a limiting growth factor in this population, although cave dwelling protects the population from predators and human disturbance.     

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.     

DIVING DEVELOPMENT AND BEHAVIOR OF A REHABILITATED MEDITERRANEAN MONK SEAL (MONACHUS MONACHUS)

Among the priority actions identified for saving the critically endangered Mediterranean monk seal are gaining basic biological information on movements and behavior, and rescuing and rehabilitating wounded, stranded, and orphaned pups. On 22 May 2004 a rehabilitated monk seal juvenile was fitted with a satellite tag, released in the National Marine Park of Alonnisos, Northern Sporades, Greece, and monitored for 167 d. Postrelease, the seal remained close to the islands of the park and within the 200-m isobath. Throughout the monitoring period, the seal reduced time hauled out, while 95-percentile dive duration and depth gradually increased. The overall maximum depth of 123 m recorded in this study is the greatest depth ever recorded for the species. These results confirm the effectiveness of the rehabilitation program carried out on the particular animal and provide additional support for the continuation of the rehabilitation program as a conservation measure for the species. We demonstrate that satellite tracking of rehabilitated seals is a valuable research and conservation tool, even for a species that commonly uses shoreline caves for resting, molting, and parturition.     

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.