Search USGSSearch

Geology topics

Mark S. Udevitz

Publications and source records attributed to Mark S. Udevitz.

At least 19 recordsLinked to original sources

Design considerations for estimating survival rates with standing age structures

Survival rate estimates are critical to understanding the dynamics and status of a population, and they are often inferred from samples of the population’s age structure. A recently developed method uses time series of standing age-structure data with information about population growth rate or fecundity to provide explicit maximum likelihood estimators of age-specific survival rates, without assuming population stability or stationarity. We explored properties of these estimators and determined sample size requirements for the estimators to achieve desired levels of precision, limit bias, and limit the probability a rate will be inestimable or its estimate inadmissible (>1).We show that estimating combined rates for adjacent age classes is an effective method of overcoming sensitivity to sampling noise in situations where collecting a larger sample of data is not feasible.

Wildlife Society Bulletin

Demography of the Pacific walrus (Odobenus rosmarus divergens) in a changing Arctic

The Pacific walrus ( Odobenus rosmarus divergens ) is a candidate to be listed as an endangered species under United States law, in part, because of climate change‐related concerns. While the population was known to be declining in the 1980s and 1990s, its recent status has not been determined. We developed Bayesian models of walrus population dynamics to assess the population by synthesizing information on population sizes, age structures, reproductive rates, and harvests for 1974–2015. Candidate models allowed for temporal variation in some or all vital rates, as well as density dependence or density independence in reproduction and calf survival. All selected models indicated that the population underwent a multidecade decline, which began moderating in the 1990s, and that annual reproductive rate and natural calf survival rates rose over time in a density‐dependent manner. However, selected models were equivocal regarding whether the natural juvenile survival rate was constant or decreasing over time. Depending on whether juvenile survival decreased after 1998, the population growth rate either increased during 1999–2015 or stabilized at a lesser level of decline than seen in the 1980s. The probability that the population was still declining in 2015 ranged from 45% to 87%.

Marine Mammal Science

Walrus haul-out and in water activity levels relative to sea ice availability in the Chukchi Sea

An animal’s energetic costs are dependent on the amount of time it allocates to various behavioral activities. For Arctic pinnipeds, the time allocated to active and resting behaviors could change with future reductions in sea ice cover and longer periods of open water. The Pacific walrus ( Odobenus rosmarus divergens ) is a large Arctic pinniped that rests on sea ice or land between foraging trips to feed on the seafloor. We used behavioral data collected from radiotagged walruses in the Chukchi Sea (2008–2014) in a Bayesian generalized linear mixed effects model to estimate the probability a walrus was in water foraging, in water not foraging, or hauled out, as a function of environmental covariates. The probability of a walrus being in water increased with wind speed and decreased with air temperature, and the probability a walrus was foraging, given it was in water, increased with available benthic macrofaunal biomass. The probability of each behavior was also related to the nature and availability of haul-out substrates. The amount of time walruses spent in water foraging and hauled out was greatest when only sea ice was available, which typically occurs when walruses occupy feeding areas during summer and early autumn. This situation may be most energy efficient for walruses because it allows the highest proportion of in water energy expenditure to be allocated to foraging. Conversely, the amount of time walruses spent in water foraging and hauled out was lowest when only land was available, which typically occurs in late autumn, in years when walruses were constrained to land haul-outs because sea ice was absent over the continental shelf.

Chukchi Sea

Evaluation of a method using survey counts and tag data to estimate the number of Pacific walruses ( Odobenus rosmarus divergens ) using a coastal haulout in northwestern Alaska

Increased periods of sparse sea ice over the continental shelf of the Chukchi Sea in late summer have reduced offshore haulout habitat for Pacific walruses ( Odobenus rosmarus divergens ) and increased opportunities for human activities in the region. Knowing how many walruses could be affected by human activities would be useful to conservation decisions. Currently, there are no adequate estimates of walrus abundance in the northeastern Chukchi Sea during summer–early autumn. Estimating abundance in autumn might be possible from coastal surveys of hauled out walruses during periods when offshore sea ice is unavailable to walruses. We evaluated methods to estimate the size of the walrus population that was using a haulout on the coast of northwestern Alaska in autumn by using aerial photography to count the number of hauled out walruses (herd size) and data from 37 tagged walruses to estimate availability (proportion of population hauled out). We used two methods to estimate availability, direct proportions of hauled out tagged walruses and smoothed proportions using local polynomial regression. Point estimates of herd size (4200–38,000 walruses) and total population size (76,000–287,000 walruses) ranged widely among days and between the two methods of estimating availability. Estimates of population size were influenced most by variation in estimates of availability. Coastal surveys might be improved most by counting walruses when the greatest numbers are hauled out, thereby reducing the influence of availability on population size estimates. The chance of collecting data during peak haulout periods would be improved by conducting multiple surveys.

Polar Biology

Sex-specific energetics of Pacific walruses ( Odobenus rosmarus divergens ) during the nursing interval

Habitat use and activity patterns of Pacific walruses ( Odobenus rosmarus divergens ) have changed with climate-induced reductions in sea ice. Increases in the time active in water could result in negative energy balance, precluding females from sustaining lactation, which could impact population demographics. Little is known about lactation costs in walruses. We examined the energetics of 0–2-yr-old walrus calves by using Bayesian hierarchical models based on longitudinal husbandry records of growth ( n = 6 females and 7 males) and caloric intake ( n = 5 females and 6 males) as a proxy for maternal lactation costs. Males and females had similar growth patterns; mean mass increased from 68 kg at birth to 301 kg by 2 yr. Females had a 2,000 kcal kg −1 higher mass storage (growth) cost than males; females typically synthesize and deposit greater amounts of adipose, which is more energy dense than lean tissue. In contrast, males had higher metabolic (basal and activity) costs, ranging from 600 to 1,800 kcal d −1 greater than similarly sized females; males are typically leaner, and muscle is more metabolically active than adipose. Yet total daily energy requirements (storage plus metabolic components) were similar across sexes, summing to approximately 190,000 kcal over the first month postpartum. Based on these estimates and assuming that 8,103 kcal is recovered from 1 kg of mass loss in adult female walruses, suckling calves could deplete 23 kg of their mother’s body mass over the first month after parturition if none of the lactation costs is met through ingested prey.

Physiological and Biochemical Zoology

Forecasting wildlife response to rapid warming in the Alaskan Arctic

Arctic wildlife species face a dynamic and increasingly novel environment because of climate warming and the associated increase in human activity. Both marine and terrestrial environments are undergoing rapid environmental shifts, including loss of sea ice, permafrost degradation, and altered biogeochemical fluxes. Forecasting wildlife responses to climate change can facilitate proactive decisions that balance stewardship with resource development. In this article, we discuss the primary and secondary responses to physical climate-related drivers in the Arctic , associated wildlife responses, and additional sources of complexity in forecasting wildlife population outcomes. Although the effects of warming on wildlife populations are becoming increasingly well documented in the scientific literature, clear mechanistic links are often difficult to establish. An integrated science approach and robust modeling tools are necessary to make predictions and determine resiliency to change. We provide a conceptual framework and introduce examples relevant for developing wildlife forecasts useful to management decisions. © 2015 Published by Oxford University Press on behalf of the American Institute of Biological Sciences 2014. This work is written by US Government employees and is in the public domain in the US.

Alaska

Identifying a reliable blubber measurement site to assess body condition in a marine mammal with topographically variable blubber, the Pacific walrus

Pacific walruses may be unable to meet caloric requirements in the changing Arctic ecosystem, which could affect body condition and have population-level consequences. Body condition has historically been monitored by measuring blubber thickness over the xiphoid process (sternum). This may be an unreliable condition index because blubber at other sites along the body may be preferentially targeted to balance energetic demands. Animals in aquaria provided an opportunity for controlled study of how blubber topography is altered by caloric intake. Morphology, body mass, blubber thickness (21 sites), and caloric intake of five mature, nonpregnant, nonlactating female walruses were measured monthly (12 month minimum). Body condition (mass × standard length −1 ) was described by a model that included caloric intake and a seasonal effect, and scaled positively with estimates of total blubber mass. Blubber thicknesses (1.91–10.69 cm) varied topographically and were similar to values reported for free-ranging female walruses. Body condition was most closely related to blubber thickness measured dorsomedially in the region of the anterior insertion of the pectoral flippers (shoulders); sternum blubber thickness was a relatively poor indicator of condition. This study demonstrates the importance of validating condition metrics before using them to monitor free-ranging populations.

Marine Mammal Science

Demography of the Pacific walrus ( Odobenus rosmarus divergens ): 1974-2006

Global climate change may fundamentally alter population dynamics of many species for which baseline population parameter estimates are imprecise or lacking. Historically, the Pacific walrus is thought to have been limited by harvest, but it may become limited by global warming-induced reductions in sea ice. Loss of sea ice, on which walruses rest between foraging bouts, may reduce access to food, thus lowering vital rates. Rigorous walrus survival rate estimates do not exist, and other population parameter estimates are out of date or have well-documented bias and imprecision. To provide useful population parameter estimates we developed a Bayesian, hidden process demographic model of walrus population dynamics from 1974 through 2006 that combined annual age-specific harvest estimates with five population size estimates, six standing age structure estimates, and two reproductive rate estimates. Median density independent natural survival was high for juveniles (0.97) and adults (0.99), and annual density dependent vital rates rose from 0.06 to 0.11 for reproduction, 0.31 to 0.59 for survival of neonatal calves, and 0.39 to 0.85 for survival of older calves, concomitant with a population decline. This integrated population model provides a baseline for estimating changing population dynamics resulting from changing harvests or sea ice.

Marine Mammal Science

Energy demands for maintenance, growth, pregnancy, and lactation of female Pacific walruses ( Odobenus rosmarus divergens )

Decreases in sea ice have altered habitat use and activity patterns of female Pacific walruses Odobenus rosmarus divergens and could affect their energetic demands, reproductive success, and population status. However, a lack of physiological data from walruses has hampered efforts to develop the bioenergetics models required for fully understanding potential population-level impacts. We analyzed long-term longitudinal data sets of caloric consumption and body mass from nine female Pacific walruses housed at six aquaria using a hierarchical Bayesian approach to quantify relative energetic demands for maintenance, growth, pregnancy, and lactation. By examining body mass fluctuations in response to food consumption, the model explicitly uncoupled caloric demand from caloric intake. This is important for pinnipeds because they sequester and deplete large quantities of lipids throughout their lifetimes. Model outputs were scaled to account for activity levels typical of free-ranging Pacific walruses, averaging 83% of the time active in water and 17% of the time hauled-out resting. Estimated caloric requirements ranged from 26,900 kcal d−1 for 2-yr-olds to 93,370 kcal d−1 for simultaneously lactating and pregnant walruses. Daily consumption requirements were higher for pregnancy than lactation, reflecting energetic demands of increasing body size and lipid deposition during pregnancy. Although walruses forage during lactation, fat sequestered during pregnancy sustained 27% of caloric requirements during the first month of lactation, suggesting that walruses use a mixed strategy of capital and income breeding. Ultimately, this model will aid in our understanding of the energetic and population consequences of sea ice loss.

Physiological and Biochemical Zoology

Estimating age ratios and size of Pacific walrus herds on coastal haulouts using video imaging

During Arctic summers, sea ice provides resting habitat for Pacific walruses as it drifts over foraging areas in the eastern Chukchi Sea. Climate-driven reductions in sea ice have recently created ice-free conditions in the Chukchi Sea by late summer causing walruses to rest at coastal haulouts along the Chukotka and Alaska coasts, which provides an opportunity to study walruses at relatively accessible locations. Walrus age can be determined from the ratio of tusk length to snout dimensions. We evaluated use of images obtained from a gyro-stabilized video system mounted on a helicopter flying at high altitudes (to avoid disturbance) to classify the sex and age of walruses hauled out on Alaska beaches in 2010–2011. We were able to classify 95% of randomly selected individuals to either an 8- or 3-category age class, and we found measurement-based age classifications were more repeatable than visual classifications when using images presenting the correct head profile. Herd density at coastal haulouts averaged 0.88 walruses/m 2 (std. err. = 0.02), herd size ranged from 8,300 to 19,400 (CV 0.03–0.06) and we documented ~30,000 animals along ~1 km of beach in 2011. Within the herds, dependent walruses (0–2 yr-olds) tended to be located closer to water, and this tendency became more pronounced as the herd spent more time on the beach. Therefore, unbiased estimation of herd age-ratios will require a sampling design that allows for spatial and temporal structuring. In addition, randomly sampling walruses available at the edge of the herd for other purposes (e.g., tagging, biopsying) will not sample walruses with an age structure representative of the herd. Sea ice losses are projected to continue, and population age structure data collected with aerial videography at coastal haulouts may provide demographic information vital to ongoing efforts to understand effects of climate change on this species.

PLoS ONE

Abundance: Population size and density estimation

Estimates of population size (total number of individuals) or density (number of individuals per unit area) are some of the most basic requirements for wildlife research and management. This article provides a brief overview of approaches for wildlife population estimation. These include habitat‐based approaches such as quadrat, line intercept, distance, and repeated count methods, as well as animal‐based approaches such as capture–recapture and harvest methods. In contrast to methods that only index changes in these parameters, estimates of size or density must account for the probability that an individual will be included in the sample used for estimation. The essential differences in the approaches are in how the samples are obtained and how the sampling probabilities are estimated.

Book chapter

Change-in-ratio

Change-in-ratio (CIR) methods are used to estimate parameters for ecological populations subject to differential removals from population subclasses. Subclasses can be defined according to criteria such as sex, age, or size of individuals. Removals are generally in the form of closely monitored sport or commercial harvests. Estimation is based on observed changes in subclass proportions caused by the removals.

Book chapter

Potential population-level effects of increased haulout-related mortality of Pacific walrus calves

Availability of summer sea ice has been decreasing in the Chukchi Sea during recent decades, and increasing numbers of Pacific walruses have begun using coastal haulouts in late summer during years when sea ice retreats beyond the continental shelf. Calves and yearlings are particularly susceptible to being crushed during disturbance events that cause the herd to panic and stampede at these large haulouts, but the potential population-level effects of this mortality are unknown. We used recent harvest data, along with previous assumptions about demographic parameters for this population, to estimate female population size and structure in 2009 and project these numbers forward using a range of assumptions about future harvests and haulout-related mortality that might result from increased use of coastal haulouts during late summer. We found that if demographic parameters were held constant, the levels of harvest that occurred during 1990–2008 would have allowed the population to grow during that period. Our projections indicate, however, that an increase in haulout-related mortality affecting only calves has a greater effect on the population than an equivalent increase in harvest-related mortality distributed among all age classes. Therefore, disturbance-related mortality of calves at coastal haulouts may have relatively important population consequences.

Polar Biology

Estimating survival rates with time series of standing age‐structure data

It has long been recognized that age‐structure data contain useful information for assessing the status and dynamics of wildlife populations. For example, age‐specific survival rates can be estimated with just a single sample from the age distribution of a stable, stationary population. For a population that is not stable, age‐specific survival rates can be estimated using techniques such as inverse methods that combine time series of age‐structure data with other demographic data. However, estimation of survival rates using these methods typically requires numerical optimization, a relatively long time series of data, and smoothing or other constraints to provide useful estimates. We developed general models for possibly unstable populations that combine time series of age‐structure data with other demographic data to provide explicit maximum likelihood estimators of age‐specific survival rates with as few as two years of data. As an example, we applied these methods to estimate survival rates for female bison ( Bison bison ) in Yellowstone National Park, USA. This approach provides a simple tool for monitoring survival rates based on age‐structure data.

Ecology

Polar bear and walrus response to the rapid decline in Arctic sea ice

The Arctic is warming faster than other regions of the world due to positive climate feedbacks associated with loss of snow and ice. One highly visible consequence has been a rapid decline in Arctic sea ice over the past 3 decades - a decline projected to continue and result in ice-free summers likely as soon as 2030. The polar bear ( Ursus maritimus ) and the Pacific walrus ( Odobenus rosmarus divergens ) are dependent on sea ice over the continental shelves of the Arctic Ocean's marginal seas. The continental shelves are shallow regions with high biological productivity, supporting abundant marine life within the water column and on the sea floor. Polar bears use sea ice as a platform for hunting ice seals; walruses use sea ice as a resting platform between dives to forage for clams and other bottom-dwelling invertebrates. How have sea ice changes affected polar bears and walruses? How will anticipated changes affect them in the future?

Alaska

Bioenergetics model for estimating food requirements of female Pacific walruses ( Odobenus rosmarus divergens )

Pacific walruses Odobenus rosmarus divergens use sea ice as a platform for resting, nursing, and accessing extensive benthic foraging grounds. The extent of summer sea ice in the Chukchi Sea has decreased substantially in recent decades, causing walruses to alter habitat use and activity patterns which could affect their energy requirements. We developed a bioenergetics model to estimate caloric demand of female walruses, accounting for maintenance, growth, activity (active in-water and hauled-out resting), molt, and reproductive costs. Estimates for non-reproductive females 0–12 yr old (65−810 kg) ranged from 16359 to 68960 kcal d −1 (74−257 kcal d −1 kg −1 ) for years with readily available sea ice for which we assumed animals spent 83% of their time in water. This translated into the energy content of 3200–5960 clams per day, equivalent to 7–8% and 14–9% of body mass per day for 5–12 and 2–4 yr olds, respectively. Estimated consumption rates of 12 yr old females were minimally affected by pregnancy, but lactation had a large impact, increasing consumption rates to 15% of body mass per day. Increasing the proportion of time in water to 93%, as might happen if walruses were required to spend more time foraging during ice-free periods, increased daily caloric demand by 6–7% for non-lactating females. We provide the first bioenergetics-based estimates of energy requirements for walruses and a first step towards establishing bioenergetic linkages between demography and prey requirements that can ultimately be used in predicting this population’s response to environmental change.

Marine Ecology Progress Series

Results and evaluation of a survey to estimate Pacific walrus population size, 2006

In spring 2006, we conducted a collaborative U.S.-Russia survey to estimate abundance of the Pacific walrus ( Odobenus rosmarus divergens ). The Bering Sea was partitioned into survey blocks, and a systematic random sample of transects within a subset of the blocks was surveyed with airborne thermal scanners using standard strip-transect methodology. Counts of walruses in photographed groups were used to model the relation between thermal signatures and the number of walruses in groups, which was used to estimate the number of walruses in groups that were detected by the scanner but not photographed. We also modeled the probability of thermally detecting various-sized walrus groups to estimate the number of walruses in groups undetected by the scanner. We used data from radio-tagged walruses to adjust on-ice estimates to account for walruses in the water during the survey. The estimated area of available habitat averaged 668,000 km 2 and the area of surveyed blocks was 318,204 km 2 . The number of Pacific walruses within the surveyed area was estimated at 129,000 with 95% confidence limits of 55,000 to 507,000 individuals. This value can be used by managers as a minimum estimate of the total population size.

Bering Sea

Divergent movements of walrus and sea ice in the northern Bering Sea

The Pacific walrus Odobenus rosmarus divergens is a large Arctic pinniped of the Chukchi and Bering Seas. Reductions of sea ice projected to occur in the Arctic by mid-century raise concerns for conservation of the Pacific walrus. To understand the significance of sea ice loss to the viability of walruses, it would be useful to better understand the spatial associations between the movements of sea ice and walruses. We investigated whether local-scale (~1 to 100 km) walrus movements correspond to movements of sea ice in the Bering Sea in early spring, using locations from radio-tracked walruses and measures of ice floe movements from processed synthetic aperture radar satellite imagery. We used generalized linear mixed-effects models to analyze the angle between walrus and ice floe movement vectors and the distance between the final geographic position of walruses and their associated ice floes (displacement), as functions of observation duration, proportion of time the walrus was in water, and geographic region. Analyses were based on 121 walrus-ice vector pairs and observations lasting 12 to 36 h. Angles and displacements increased with observation duration, proportion of time the walrus spent in the water, and varied among regions (regional mean angles ranged from 40° to 81° and mean displacements ranged from 15 to 35 km). Our results indicated a lack of correspondence between walruses and their initially associated ice floes, suggesting that local areas of walrus activities were independent of the movement of ice floes.

Marine Ecology Progress Series