Non-breeding changes in at-sea distribution and abundance of the threatened marbled murrelet (Brachyramphus marmoratus) in a portion of its range exhibiting long-term breeding season declines.

The marbled murrelet (Brachyramphus marmoratus) is classified as a threatened species under the US Endangered Species Act in Washington, Oregon, and California USA due to population declines, loss of breeding habitat, and other factors. To date, population assessments have focused on breeding season at-sea surveys. Consequently, there is little information on this species' distribution, abundance, and population trends during the non-breeding season, when murrelets are found exclusively in the marine environment. To address this information need, we assessed non-breeding (Sep-Mar) at-sea murrelet abundance patterns and population trends over 8 years, in a portion of its range where breeding season surveys indicate a 20-year population decline, Puget Sound, Washington, USA. This allowed us to assess whether non-breeding population trends mirrored those observed during the breeding season suggesting regional year-round conservation concerns and to also identify important over-wintering areas (areas of high abundance). We integrated our non-breeding abundance information with breeding season information to assess year-round patterns of abundance. This allowed us to test the prediction that murrelets move into the relatively protected inner marine waters of Puget Sound from harsher outer coastal habitats during the non-breeding season to molt and over-winter. Similar to trends from the breeding season, we observed strong murrelet density declines across the entire non-breeding period (Sep and Apr) with declines most pronounced in the fall and early winter (lateSep-Dec) survey windows when birds molt and in the spring just prior to breeding (Mar-Apr). Despite these declines, there was essentially no change in murrelet density in mid-winter (January-February) when overall density was lower. Puget Sound murrelet density exhibited a strong north-south gradient with relatively high densities to the north and low densities to the south; murrelets were largely absent from Central Puget Sound. For strata other than Central Puget Sound, density varied seasonally with birds more evenly distributed among strata between September and December but in the late winter/early spring period (Jan-Apr), murrelets were largely absent from all strata except the most northerly Admiralty Inlet Stratum, which appears to be important to murrelets year-round. Depending on the year, non-breeding season densities were nearly the same or higher than breeding season densities indicate that murrelets were not moving into the relatively protected inner marine waters of Puget Sound from more outer coastal environments during the non-breeding season as predicted.

Conservation planning for species recovery under the Endangered Species Act: A case study with the Northern Spotted Owl.

The northern spotted owl (Strix occidentalis caurina) was listed as threatened under the U.S. Endangered Species Act (ESA) in 1990. We applied modern spatial conservation theory and models to evaluate several candidate critical habitat networks, and sought an efficient conservation solution that encompassed the highest value lands for spotted owl recovery rather than maximizing the total area of potential critical habitat. We created a map of relative habitat suitability, which served as input to the spatial conservation prioritization program Zonation. We used the spatially-explicit individual-based population model HexSim to estimate and compare simulated spotted owl population outcomes among a suite of candidate critical habitat networks that varied in size and spatial arrangement under alternative scenarios of future habitat suitability and barred owl (S. varia) effects. We evaluated simulated spotted owl population outcomes, including total population size, and extinction and quasi-extinction likelihoods for 108 combinations of candidate critical habitat networks by habitat change by barred owl scenarios, both range-wide and within 11 distinct portions of the owl's range. Barred owl encounter rates and the amount and suitability of habitat had substantial effects on simulated spotted owl populations. When barred owl encounter rates were high, changes in the amount and suitability of habitat had minimal impacts on population performance. Under lowered barred owl encounter rates, candidate critical habitat networks that included most existing high suitability habitat supported a high likelihood of long-term population persistence. Barred owls are currently the primary driving force behind poor population performance of NSOs; however, our models demonstrated that a sufficient area of high suitability habitat remains essential for recovery when effects of barred owls can be reduced. The modeling approach we employed is sufficiently flexible to incorporate new information about spotted owls as it becomes available and could likely be applied to conservation planning for other species.

Marine Habitat Selection by Marbled Murrelets (Brachyramphus marmoratus) during the Breeding Season.

The marbled murrelet (Brachyramphus marmoratus) is a declining seabird that is well-known for nesting in coastal old-growth forests in the Pacific Northwest. Most studies of habitat selection have focused on modeling terrestrial nesting habitat even though marine habitat is believed to be a major contributor to population declines in some regions. To address this information gap, we conducted a 5-year study of marine resource selection by murrelets in Washington, which contains a population experiencing the steepest documented declines and where marine habitat is believed to be compromised. Across five years we tracked 157 radio-tagged murrelets during the breeding season (May to August), and used discrete choice models to examine habitat selection. Using an information theoretic approach, our global model had the most support, suggesting that murrelet resource selection at-sea is affected by many factors, both terrestrial and marine. Locations with higher amounts of nesting habitat (ß = 21.49, P < 0.001) that were closer to shore (ß = -0.0007, P < 0.001) and in cool waters (ß = -0.2026, P < 0.001) with low footprint (ß = -0.0087, P < 0.001) had higher probabilities of use. While past conservation efforts have focused on protecting terrestrial nesting habitat, we echo many past studies calling for future efforts to protect marine habitat for murrelets, as the current emphasis on terrestrial habitat alone may be insufficient for conserving populations. In particular, marine areas in close proximity to old-growth nesting habitat appear important for murrelets during the breeding season and should be priorities for protection.

Assessing ecological correlates of marine bird declines to inform marine conservation.

Identifying drivers of ecosystem change in large marine ecosystems is central for their effective management and conservation. This is a sizable challenge, particularly in ecosystems transcending international borders, where monitoring and conservation of long-range migratory species and their habitats are logistically and financially problematic. Here, using tools borrowed from epidemiology, we elucidated common drivers underlying species declines within a marine ecosystem, much in the way epidemiological analyses evaluate risk factors for negative health outcomes to better inform decisions. Thus, we identified ecological traits and dietary specializations associated with species declines in a community of marine predators that could be reflective of ecosystem change. To do so, we integrated count data from winter surveys collected in long-term marine bird monitoring programs conducted throughout the Salish Sea--a transboundary large marine ecosystem in North America's Pacific Northwest. We found that decadal declines in winter counts were most prevalent among pursuit divers such as alcids (Alcidae) and grebes (Podicipedidae) that have specialized diets based on forage fish, and that wide-ranging species without local breeding colonies were more prone to these declines. Although a combination of factors is most likely driving declines of diving forage fish specialists, we propose that changes in the availability of low-trophic prey may be forcing wintering range shifts of diving birds in the Salish Sea. Such a synthesis of long-term trends in a marine predator community not only provides unique insights into the types of species that are at risk of extirpation and why, but may also inform proactive conservation measures to counteract threats--information that is paramount for species-specific and ecosystem-wide conservation.

Spatiotemporal variation in resource selection: insights from the American marten (Martes americana).

Behavioral and genetic adaptations to spatiotemporal variation in habitat conditions allow species to maximize their biogeographic range and persist over time in dynamic environments. An understanding of these local adaptations can be used to guide management and conservation of populations over broad extents encompassing diverse habitats. This understanding is often achieved by identifying covariates related to species' occurrence in multiple independent studies conducted in relevant habitats and seasons. However, synthesis across studies is made difficult by differences in the model covariates evaluated and analytical frameworks employed. Furthermore, inferences may be confounded by spatiotemporal variation in which habitat attributes are limiting to the species' ecological requirements. In this study, we sought to quantify spatiotemporal variation in resource selection by the American marten (Martes americana) in forest ecosystems of the Pacific Northwest, USA. We developed resource selection functions for both summer and winter based on occurrence data collected in mesic and xeric forest habitats. Use of a consistent analytical framework facilitated comparisons. Habitat attributes predicting marten occurrence differed strongly between the two study areas, but not between seasons. Moreover, the spatial scale over which covariates were calculated greatly influenced their predictive power. In the mesic environment, marten resource selection was strongly tied to riparian habitats, whereas in the xeric environment, marten responded primarily to canopy cover and forest fragmentation. These differences in covariates associated with marten occurrence reflect differences in which factors were limiting to marten ecology in each study area, as well as local adaptations to habitat variability. Our results highlight the benefit of controlled meta-replication studies in which analyses of multiple study areas and seasons at varying spatial scales are integrated into a single framework.

Genetic analyses of historic and modern marbled murrelets suggest decoupling of migration and gene flow after habitat fragmentation.

The dispersal of individuals among fragmented populations is generally thought to prevent genetic and demographic isolation, and ultimately reduce extinction risk. In this study, we show that a century of reduction in coastal old-growth forests, as well as a number of other environmental factors, has probably resulted in the genetic divergence of marbled murrelets (Brachyramphus marmoratus) in central California, despite the fact that 7 per cent of modern-sampled murrelets in this population were classified as migrants using genetic assignment tests. Genetic differentiation appears to persist because individuals dispersing from northern populations contributed relatively few young to the central California population, as indicated by the fact that migrants were much less likely to be members of parent-offspring pairs than residents (10.5% versus 45.4%). Moreover, a recent 1.4 per cent annual increase in the proportion of migrants in central California, without appreciable reproduction, may have masked an underlying decline in the resident population without resulting in demographic rescue. Our results emphasize the need to understand the behaviour of migrants and the extent to which they contribute offspring in order to determine whether dispersal results in gene flow and prevents declines in resident populations.

Characterizing dispersal patterns in a threatened seabird with limited genetic structure.

Genetic assignment methods provide an appealing approach for characterizing dispersal patterns on ecological time scales, but require sufficient genetic differentiation to accurately identify migrants and a large enough sample size of migrants to, for example, compare dispersal between sexes or age classes. We demonstrate that assignment methods can be rigorously used to characterize dispersal patterns in a marbled murrelet (Brachyramphus marmoratus) population from central California that numbers approximately 600 individuals and is only moderately differentiated (F(ST) approximately 0.03) from larger populations to the north. We used coalescent simulations to select a significance level that resulted in a low and approximately equal expected number of type I and II errors and then used this significance level to identify a population of origin for 589 individuals genotyped at 13 microsatellite loci. The proportion of migrants in central California was greatest during winter when 83% of individuals were classified as migrants compared to lower proportions during the breeding (6%) and post-breeding (8%) seasons. Dispersal was also biased toward young and female individuals, as is typical in birds. Migrants were rarely members of parent-offspring pairs, suggesting that they contributed few young to the central California population. A greater number of migrants than expected under equilibrium conditions, a lack of individuals with mixed ancestry, and a small number of potential source populations (two), likely allowed us to use assignment methods to rigorously characterize dispersal patterns for a population that was larger and less differentiated than typically thought required for the identification of migrants.

Conservation of the marbled murrelet under the Northwest Forest Plan.

The Marbled Murrelet (Brachyramphus marmoratus) was listed as threatened in 1992, primarily because of loss of its old-forest nesting habitat. Monitoring conducted over the first 10 years following implementation of the Northwest Forest Plan shows at-sea murrelet populations appear to be stationary, but recruitment is very low and demographic models project a 4-6% annual rate of decline. Monitoring of nesting habitat indicated there were about 1.6 million ha of higher-suitability nesting habitat on all lands at the start of the plan, about half of which occurred on federal lands. Most (88%) of higher-suitability habitat on federal lands was protected within reserves. Over the past 10 years, losses of habitat due primarily to fire have totaled about 2% on federal lands. Losses have been much greater (12%) on nonfederal lands, due primarily to timber harvest. Habitat is expected to accrue within reserves as younger forest matures and attains sufficient diameter to support nesting sites. At-sea estimates of population size are strongly and positively correlated with amounts of adjacent nesting habitat at a broad scale, supporting the idea that amounts of nesting habitat are a primary driver in wide-scale murrelet population distribution. Conditions at sea, however such as temperature regimes, prey availability, and pollutants, continue to affect murrelet populations. The system of large reserves seems to have achieved the short-term objective of conserving much of the remaining nesting habitat on federal lands. These reserves are also likely to contribute to the long-term objective of creating large, contiguous blocks of nesting habitat. The plan has a primary role in conserving and restoring nesting habitat on federal land but will succeed in this role only if land allocations calling for such protection are in place for many decades.

Modeling the Dynamics of Snags.

Many wildlife species required standing dead trees (i.e., snags) as part of their habitat. Therefore, the ability to predict future density, distribution, and condition of snags can assist resource managers in making land-use decisions. Here we present methods for modeling the dynamics of snags using data from a 10-yr study on the rates of decay, falling, and recruitment of snags on burned and unburned plots in the Sierra Nevada, California. Snags (all species) in advanced stages of decay usually fell within 5 yr, and snags created by fire decayed rapidly and fell quicker (within 10 yr) than those on unburned plots. Pine (Pinus spp.) snags decayed more rapidly than fir (Abies spp.). Although there was an overall net increase in snag density on unburned plots, most of this increase was in the smaller (>13-38 cm diameter at breast height [dbh]) size classes; there was a net decrease in the larger (>38 cm dbh) snags preferred by many birds for nesting and feeding. Overall, snags remained standing the longest that were larger in diameter, shorter in height, less decayed, fir rather than pine, and lacking tops. A Leslie matrix model of snag dynamics predicted changes in snag decay and density only when adjusted for the specific environmental factors(s) causing initial tree mortality. Many snags are created by episodic events, such as fire, disease, drought, and insects. Models of snag dynamics must include the species and condition of trees becoming snags, as well as the factor(s) causing the tree to die. Forest managers must consider this episodic creation of snags when developing snag-management guidelines, and when planning tree-salvage programs based on short-term inventories.