A STAGE-BASED MODEL OF MANATEE POPULATION DYNAMICS

A stage-structured population model for the Florida manatee ( Trichechus manatus latirostris ) was developed that explicitly incorporates uncertainty in parameter estimates. The growth rates calculated with this model reflect the status of the regional populations over the most recent 10-yr period. The Northwest and Upper St. Johns River regions have growth rates (λ) of 1.037 (95% interval, 1.016–1.056) and 1.062 (1.037–1.081), respectively. The Southwest region has a growth rate of 0.989 (0.946–1.024), suggesting this population has been declining at about 1.1% per year. The estimated growth rate in the Atlantic region is 1.010 (0.988–1.029), but there is some uncertainty about whether adult survival rates have been constant over the last 10 yr; using the mean survival rates from the most recent 5-yr period, the estimated growth rate in this region is 0.970 (0.938–0.998). Elasticity analysis indicates that the most effective management actions should seek to increase adult survival rates. Decomposition of the uncertainty in the growth rates indicates that uncertainty about population status can best be reduced through increased monitoring of adult survival rate.     

A population estimate of Heaviside's dolphins, Cephalorhynchus heavisidii, at the southern end of their range

Heaviside's dolphins, Cephalorhynchus heavisidii , are endemic to southwestern Africa, where they are exposed to unknown levels of anthropogenic threats, including inshore set netting. Using photo-ID data collected over 3 yr on the west coast of South Africa, we calculated Chapman's-modified Petersen estimates of the number of distinctive individuals at three spatial scales. Sample sizes were small and recapture rates low resulting in high variance. Total population abundance was extrapolated from the proportion of well-marked animals in the population (14%–17%) with between-year estimates adjusted for mortality using data from Commerson's dolphin. The total population size was calculated as 527 animals (CV = 0.35, CI 272–1,020) in the 1999 study area (20 km of coastline, within season), 3,429 animals (CV = 0.36, CI 1,721–6,828) in the central study area (150 km of coastline, 3 yr), and 6,345 animals (CV = 0.26, CI 3,573–11,267) in the full study area (390 km of coastline, 2 yr). Dolphins fitted with satellite transmitters varied in their use of the inshore photographic study area from 39.5% to 94.7% of transmission days (38–51 total). Given the known or suspected biases in the data, these abundance estimates are likely to be biased downward.     

The propagation of uncertainty in human mortality processes operating in stochastic environments

This paper presents a model describing how the uncertainty due to influential exogenous processes combines with stochasticity intrinsic to physiological aging processes and propagates through time to generate uncertainty about the future physiological state of the population. Variance expressions are derived for (a) the future values of the physiological variables under the assumption that external factors evolve under a linear stochastic diffusion process, and (b) the cohort survival functions and cohort life expectancies which reflect the uncertainty in the future values of the physiological variables. The model implies that a major component of uncertainty in forecasts of the physiological characteristics of a closed cohort is due to differential rates of survival associated with different realizations of the external process. This suggests that the limits to forecasting may be different in physiological systems subject to systematic mortality than in physical systems such as weather where the concepts of closed cohorts and of mortality selection have no simple analog.     

Mortality in Svalbard reindeer

Mortality rates in Svalbard reindeer followed the "U"-shaped pattern, with higher mortality rate among calves and old animals than in middle aged individuals. Assuming a stable population size, the mortality data predict a 16.2% annual mortality and recruitment rate among 6 months and older animals, and a male:female sex ratio of 48.5:51.5 among 1-yr old and older. Both predictions are supported by field data. Female mortality rate increased and differed from the male rate in the age interval 2-4 yr, while the male rate increased sharply and differed permanently from the female rate in the age interval above 6 yr. First and last breeding are thought to cause the increased mortality among 2–4 and 11–13 yr-old females. Increased involvement in rutting activities with age associated with lower grazing activity and depletion of fat reserves probably cause the increasing mortality rate among the 6+ yr males. Maximum life span was 17 yr in females and 12 yr in males. Apart from the unknown but possibly high neonatal calf-mortality, only few animals died during summer or during the rut in autumn. Death among calves, yearlings and most of the females occurred before antler-shedding in May-June. Most of the 2-yr and older males died after antler shedding, which occurs from December among the oldest and during April-May among the younger males. Hence, the major part of the mortality takes place in spring. The lack of visible subcutaneous and femur fat in most carcasses indicated starvation as the major mortality cause.     

Modeling population viability of captive elephants in Myanmar (Burma): implications for wild populations

Captive Asian elephants Elephas maximus , used as work animals, constitute up to 22–30% of remaining Asian elephants. Myanmar has the largest captive population worldwide (∼6000), maintained at this level for over a century. We used published demographic data to assess the viability of this captive population. We tested how this population can be self-sustained, how many elephants must be supplemented from the wild to maintain it, and what consequences live capture may have for Myanmar's wild population. Our results demonstrate that the current captive population is not self-sustaining because mortality is too high and birth rates are too low. Our models also suggest ∼100 elephants year⁻¹ have been captured in the wild to supplement the captive population. Such supplementation cannot be supported by a wild population of fewer than 4000 elephants. Given the most recent expert estimate of ∼2000 wild elephants remaining in Myanmar, a harvest of 100 elephants year⁻¹ could result in extinction of the wild population in 31 years. Continued live capture threatens the survival of wild and captive populations and must stop. In addition, captive breeding should be increased. These measures are essential to slow the decline and extinction of all of Myanmar's elephants.     

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.     

The energy balance of the Australian brine shrimp, Parartemia zietziana (Crustacea : Anostraca)

SUMMARY. Assimilation budgets (i.e. assimilation = respiration + excretion + production) are presented for cohorts of P. zietziana in two salt lakes. Shrimps in Pink Lake had an assimilation rate of 1631.6 kJ m⁻² year ⁻¹, those in Lake Cundare 212.1 kJ m⁻² year⁻¹. In both lakes, respiration accounted for 60–80% of assimilation. Assimilation rates for individuals (derived as assimilation = ingestion minus faecal output) are also given and compared with respiratory rates of individuals. The comparisons indicated that energy was often consumed at a higher rate by respiration than it could be supplied by assimilation. Starvation due to a low assimilation efficiency was suggested as a cause of the consistent mortality, variable growth rate of individuals and unpredictable recruitment which were characteristic of the cohorts of P. zietziana in both lakes. An analysis of mortality showed that the young had the poorest survival, as predicted by a theoretical model of a starving zooplankter and a comparison of the increase with weight of ingestion and respiration. Gross growth efficiency (production: assimilation) was 15–30%, about the same as published data on Anostracans. Net growth efficiency (production : consumption) was 5–12% and generally lower than published values reflecting the difficulty P. zietziana has in balancing its budget.     

Source populations in carnivore management: cougar demography and emigration in a lightly hunted population

Wildlife agencies typically attempt to manage carnivore numbers in localized game management units through hunting, and do not always consider the potential influences of immigration and emigration on the outcome of those hunting practices. However, such a closed population structure may not be an appropriate model for management of carnivore populations where immigration and emigration are important population parameters. The closed population hypothesis predicts that high hunting mortality will reduce numbers and densities of carnivores and that low hunting mortality will increase numbers and densities. By contrast, the open population hypothesis predicts that high hunting mortality may not reduce carnivore densities because of compensatory immigration, and low hunting mortality may not result in more carnivores because of compensatory emigration. Previous research supported the open population hypothesis with high immigration rates in a heavily hunted (hunting mortality rate=0.24) cougar population in northern Washington. We test the open population hypothesis and high emigration rates in a lightly hunted (hunting mortality rate=0.11) cougar population in central Washington by monitoring demography from 2002 to 2007. We used a dual sex survival/fecundity Leslie matrix to estimate closed population growth and annual census counts to estimate open population growth. The observed open population growth rate of 0.98 was lower than the closed survival/fecundity growth rates of 1.13 (deterministic) and 1.10 (stochastic), and suggests a 12–15% annual emigration rate. Our data support the open population hypothesis for lightly hunted populations of carnivores. Low hunting mortality did not result in increased numbers and densities of cougars, as commonly believed because of compensatory emigration.     

PHYSIOLOGICAL AND BEHAVIORAL DEVELOPMENT IN DELPHINID CALVES: IMPLICATIONS FOR CALF SEPARATION AND MORTALITY DUE TO TUNA PURSE-SEINE SETS

Tuna purse-seiners in the eastern tropical Pacific (ETP) capture yellowfin tuna by chasing and encircling herds of associated dolphins. This fishery has caused mortality in 14 dolphin species (20 stocks) and has led to significant depletions of at least three stocks. Although observed dolphin mortality is currently low, set frequency remains high and dolphin stocks are not recovering at expected rates. Mortality of nursing calves permanently separated from their mothers during fishery operations may be an important factor in the lack of population recovery, based on the recent discovery that calves do not accompany 75%–95% of lactating females killed in the purse-seine nets. We assessed age-specific potential for mother–calf separations and subsequent mortality of calves by reviewing and synthesizing published data on physiological and behavioral development in delphinids from birth through 3 yr postpartum. Results indicate that evasive behavior of mothers, coupled with the developmental state of calves, provides a plausible mechanism for set-related mother–calf separations and subsequent mortality of calves. Potential for set-related separation and subsequent mortality is highest for 0–12-mo-old dolphins and becomes progressively lower with age as immature dolphins approach adult stamina and attain independence.     

THE POPULATION DYNAMICS OF NORTHERN SEA LIONS, 1975-1985

Abstract: Populations of northern sea lions ( Eumetopias jubatus ) in the vicinity of Marmot Island, Alaska declined during 1975–1985 at about 5% per year (Merrick et al. 1987). The cause of this decline is not known. A life table for the northern sea lion was calculated assuming that life spans follow a Weibull distribution. Samples of northern sea lions taken in the vicinity of Marmot Island, Alaska during 1975–1978 and 1985–1986 indicate that the average age of females older than 3 yr increased about 1.55 yr (SD = 0.35 yr) while the population was declining at about 5% per year. Fecundity rates decreased by 10% over the same period, but the decrease was not statistically significant (Calkins and Goodwin 1988). Possible causes of the population decline and the change in age structure were examined by writing the Leslie matrix population equation in terms of changes in juvenile and adult survival rates and fecundity, and examining the short–term behavior of the trajectories of the average age of adult females, total number of females, and total number of pups with respect to those changes in the vital parameters. From the observed rate of declines of adults and the changes in average age of adult females and fecundity, estimates of the changes in adult and juvenile survival were calculated; estimates of the standard deviations of these changes were estimated via a bootstrap procedure. One purpose of this exercise is to aid in setting priorities for research for determining the cause of the decline. An explanation for the observed declines in numbers of adult sea lions consistent with the observed fecundity rates, a rate of decrease of 5% in the number of adults, and the corresponding increase in average age (of females age 3 yr and older) was a 10%–20% decrease in the survival of juveniles (age 0-3 yr) coupled with an insignificant change in adult survival (0.03%, SD = 1%).     

Lion density and population structure in the Selous Game Reserve: evaluation of hunting quotas and offtake

In 1992, tourist hunting in the Selous Game Reserve generated 1.28 million dollars for the Tanzanian government, of which 0·96 million dollars were returned to wildlife conservation. Lions ( Panthera leo ) are one of three critical species for tourist hunting, consistently generating 12%–13% of hunting revenue from 1988 to 1992. Because of their ecological and economic importance (and intrinsic value), it is important that lion quotas be set so that offtake is sustainable. The population density of lions in Selous ranges from 0·08 to 0·13 adults km⁻², comparable to unhunted ecosystems. The adult sex ratio (36–41% male) and the ratio of cubs to adults (29% cubs) are similar to those of unhunted populations. The ratio of lions to hyaenas is lower in heavily hunted areas (0·17 lions/hyena) than in unhunted areas (0·43 lions/hyena). Hunting levels between 1989 and 1994 took 2·7–4·3% of adult males annually, which is sustainable. The current quota is 10–16% of the adult male population, which exceeds natural mortality rates for male lions. To remain stable if the quota was filled, the population would have to compensate via increased fecundity, increased juvenile survival, or an altered sex‐ratio. Compensation occurs in Selous by producing (or raising) more male than female cubs (66–81% of juveniles are male). Only 28% of the Selous quota was filled in 1992. The percentage of quota filled (both in Selous and nationwide) has dropped since 1988 as quotas have increased. The current intensity of lion hunting in Selous is sustainable, but the quota cannot be filled sustainably.     

Population growth rates of reef sharks with and without fishing on the great barrier reef: robust estimation with multiple models

Overfishing of sharks is a global concern, with increasing numbers of species threatened by overfishing. For many sharks, both catch rates and underwater visual surveys have been criticized as indices of abundance. In this context, estimation of population trends using individual demographic rates provides an important alternative means of assessing population status. However, such estimates involve uncertainties that must be appropriately characterized to credibly and effectively inform conservation efforts and management. Incorporating uncertainties into population assessment is especially important when key demographic rates are obtained via indirect methods, as is often the case for mortality rates of marine organisms subject to fishing. Here, focusing on two reef shark species on the Great Barrier Reef, Australia, we estimated natural and total mortality rates using several indirect methods, and determined the population growth rates resulting from each. We used bootstrapping to quantify the uncertainty associated with each estimate, and to evaluate the extent of agreement between estimates. Multiple models produced highly concordant natural and total mortality rates, and associated population growth rates, once the uncertainties associated with the individual estimates were taken into account. Consensus estimates of natural and total population growth across multiple models support the hypothesis that these species are declining rapidly due to fishing, in contrast to conclusions previously drawn from catch rate trends. Moreover, quantitative projections of abundance differences on fished versus unfished reefs, based on the population growth rate estimates, are comparable to those found in previous studies using underwater visual surveys. These findings appear to justify management actions to substantially reduce the fishing mortality of reef sharks. They also highlight the potential utility of rigorously characterizing uncertainty, and applying multiple assessment methods, to obtain robust estimates of population trends in species threatened by overfishing.     

When being alive implies being safe: variation in mortality rates can cause oviposition selectivity to increase with age

Optimal behavioural decisions are expected to depend on various state variables, such as physiological condition or age. In insects, empirical evidence of the effect of adult age on oviposition selectivity is mixed. Consistently, optimality models – which primarily incorporate the effects of egg load and senescence – fail to provide universal predictions. Here we propose that spatial variation in mortality rates creates an additional mechanism able to select for an increase in selectivity with age. Females can be selected to use the fact of having reached an advanced age as a cue of low mortality rates in their environment. Older females may thus be less time-limited, and can afford for more careful host selection. This is because variation in mortality rates can cause a positive correlation between individual age, and expected residual life span. We present a simulation model that formalises the verbal argument presented above, and discuss the findings in the more general context of the dependence of reproductive output on age.     

CALCULATING LIMITS TO THE ALLOWABLE HUMAN-CAUSED MORTALITY OF CETACEANS AND PINNIPEDS

A simulation method was developed for identifying populations with levels of human-caused mortality that could lead to depletion, taking into account the uncertainty of available information. A mortality limit (termed the Potential Biological Removal, PBR , under the U. S. Marine Mammal Protection Act) was calculated as the product of a minimum population estimate ( N _MIN), one-half of the maximum net productivity rate ( R _MAX), and a recovery factor ( F _R). Mortality limits were evaluated based on whether at least 95% of the simulated populations met two criteria: (1) that populations starting at the maximum net productivity level ( MNPL ) stayed there or above after 20 yr, and (2) that populations starting at 30% of carrying-capacity ( K ) recovered to at least MNPL after 100 yr. Simulations of populations that experienced mortality equal to the PBR indicated that using approximately the 20th percentile (the lower 60% log-normal confidence limit) of the abundance estimate for N _MIN met the criteria for both cetaceans (assuming R _MAX= 0.04) and pinnipeds (assuming R _MAX= 0.12). Additional simulations that included plausible levels of bias in the available information indicated that using a value of 0.5 for F _R would meet both criteria during these "bias trials." It is concluded that any marine mammal population with an estimate of human-caused mortality that is greater than its PBR has a level of mortality that could lead to the depletion of the population. The simulation methods were also used to show how mortality limits could be calculated to meet conservation goals other than the U. S. goal of maintaining populations above MNPL .     

Life cycle, growth and production in estuarine populations of the polychaetes Nereis virens and N. diversicolor

Life cycle, growth, mortality and production of the polychaetes Nereis virens and N. diversicolor were studied in Norsminde Fjord, Denmark. When 3 yr old, N. virens showed synchronous spawning induced at new moon in April when water temperature was 10–12°C. Nereis diversicolor showed a prolonged spawning during early spring and summer at an age of 12–18 months. Average weight-specific growth rate for N. virens and N. diversicolor were O.0062 and 0.0050 d⁻¹, respectively; showing a reduced rate with age. Annual mortalities of 76–77% and 98%, respectively, reduced N. virens to 1.2–1.4% and N. diversicolor to 1.2–2.3% of the initial population at the time of spawning. Annual production of N. virens and N. diversicolor was 23.73 and 27.17 g AFDW m⁻², tantamount to an annual turnover (P/ B ) of 2.52 and 2.60, respectively.     

Comparison of discriminant function and classification tree analyses for age classification of marmots

We evaluated the predictive power of two classification techniques, one parametric – discriminant function analysis (DFA) and the other non-parametric – classification and regression tree analysis (CART), in order to provide a non-subjective quantitative method of determining age class in Vancouver Island marmots ( Marmota vancouverensis ) and hoary marmots ( Marmota caligata ). For both techniques we used morphological measurements of known-age male and female marmots from two independent population studies to build and test predictive models of age class. Both techniques had high predictive power (69–86%) for both sexes and both species. Overall, the two methods performed identically with 81% correct classification. DFA was marginally better at discriminating among older more challenging age classes compared to CART. However, in our test samples, cases with missing values in any of the discriminant variables were deleted and hence unclassified by DFA, whereas CART used values from closely correlated variables to substitute for the missing values. Therefore, overall, CART performed better (CART 81% vs DFA 76%) because of its ability to classify incomplete cases. Correct classification rates were approximately 10% higher for hoary marmots than for Vancouver Island marmots, a result that could be attributed to different sets of morphological measurements. Zygomatic arch breadth measured in hoary marmots was the most important predictor of age class in both sexes using both classification techniques. We recommend that CART analysis be performed on data-sets with incomplete records and used as a variable screening tool prior to DFA on more complete data-sets.     

False killer whales (Pseudorca crassidens) around the main Hawaiian Islands: Long-term site fidelity, inter-island movements, and association patterns

Despite their world-wide distribution throughout the tropics and subtropics, false killer whales ( Pseudorca crassidens ) are one of the lesser-known large odontocetes. Genetic evidence indicates a demographically isolated population around the main Hawaiian Islands. We examine site fidelity, movements and association patterns in this population using data from directed surveys and opportunistic photographs from 1986 to 2007. This species was only infrequently encountered, and while found in depths from 38 to 4,331 m, sighting rates were greatest in depths >3,000 m. We photo-identified 152 distinctive individuals. Resighting rates were high, with an average of 76.8% of distinctive individuals within groups documented on more than one occasion. Most (86.6%) were linked by association into a single social network; only one large group (16 distinctive individuals), documented the farthest offshore (42–70 km), did not link by association to that large network, and may be part of an offshore population. Individual movements of up to 283 km were documented, with a large proportion of individuals moving among islands. Individuals were resighted up to 20.1 yr after first being documented, showing long-term fidelity to the islands. Repeated associations among individuals were documented for up to 15 yr, and association analyses indicate preferred associations and strong bonds among individuals.     

THE LIFE HISTORY OF FREE-RANGING ATLANTIC SPOTTED DOLPHINS (STENELLA FRONTALIS): AGE CLASSES, COLOR PHASES, AND FEMALE REPRODUCTION

Atlantic spotted dolphins (Stenella frontalis) were observed underwater and from the surface from 1985 to 1996 and photographed through successive years. Individuals were categorized into age classes by their degree of spotting and color phases. Dolphins spent an average of 3 yr in the two-tone color phase, 5 yr in the speckled phase, 7 yr in the mottled phase and up to 10 yr or more in the fused phase.
Sex ratios were close to parity, with old adults skewed towards females and juveniles and young adults skewed towards males. The average calving interval for 24 females was 2.96 years with a range of 1–5 yr. Females whose calves survived the first year had a significantly longer calving interval (3.56 years). The ages of first parturition for five females were estimated to be 10–12 yr. The age at sexual maturation was estimated to range from 8 to 15 yr.
Pregnancy rate fluctuated annually, with an average rate of 0.25 (range 0.07–0.57). Annual average birth rate was 0.08 (range 0.07–0.14), average calf production was 0.33 (range 0.06–0.52), average fecundity was 0.23 (range 0.13–0.30), and average recruitment was 0.06 (range 0.03–0.08). Most females who lost a calf conceived the same or following year.
Lactation lasted up to 5 yr, and 45% of visibly pregnant females were also lactating. Age of first parturition was associated with the mottled color phase. Average first-year mortality rate of calves was 0.24.     

The prevalence of non-breeders in raptor populations: evidence from rings, radio-tags and transect surveys

Age-specific survival and breeding (ASSAB) models were developed with data from 146 common buzzards ( Buteo buteo ) radio-tagged in southern Britain during 1990–1998, in a 120-km2 study area that had on average 25 egg-laying pairs. Survival checks were aided by philopatric behaviour and a maximum annual tag failure rate of 7%: minimum survival rates, that were estimated by assuming death of buzzards with lost tags, were close to maximum rates that were estimated using only the recorded deaths. First-year survival rate estimates for 35 buzzards fitted in 1990–1991 with 25–30-g backpack radios were 69–74% (minimum-maximum), close to the 61–71% for 16 buzzards with 12-g tail-mount radios; the backpacks transmitted for 2–4 yr. Overall survival rates were 66–73% in the first year, 91–97% in the second and 88–91% thereafter. Survival estimates from 288 recent British ring recoveries were lower in the first and second years, at 55% and 75%, but similar (88%) thereafter. Most deaths were from natural causes (40%) or interaction with artefacts (36%). ASSAB models, from radio-tracking and the observed 1.71 young clutch−1, predicted breeding by 16–21% of all the buzzards present in spring, or up to 25% with the minimum likely productivity of 1.4 young clutch−1 or 12% net emigration. Ringing data predicted breeding rates of 33–38%. The models were tested with density data from nest surveys and new radio-corrected-transect and truncation-mark-resighting estimates of buzzard numbers. Surveys in autumn and late winter estimated breeding rates of 21–25%. The high non-breeder density in spring, of three other buzzards for each paired bird with eggs, has important implications for understanding evolutionary fitness, predation and population ecology.