Scott M. Gende

Pacific Salmon in Aquatic and Terrestrial Ecosystems

S runs in the Pacific Northwest have been declining for decades, so much so that many runs are threatened or endangered; others have been completely extirpated (Nehlsen et al. 1991). This “salmon crisis” looms large in the public eye, because it has serious and wideranging economic, cultural, and ecological repercussions. Billions of dollars have gone into industrial and agricultural projects that alter regional rivers in ways that, often unintentionally, make them inaccessible or unsuitable for salmon. Recently, billions more have been spent in largely unsuccessful attempts to restore the languishing salmon runs (Lichatowich 1999). Moreover, enormous nonmonetary resources have been expended in assigning and denying responsibility for failed runs and debating the possible efficacy of various remedies. As resources that are devoted to reversing declining runs of salmon have increased, scientists and resource managers have been expanding our understanding of the ecological role of salmon and other anadromous fishes, which return from the sea to spawn in fresh water. We have known for years that spawning salmon serve as a food resource for wildlife species (e.g., Shuman 1950) and, when they die after spawning (as most Pacific salmon do), their carcasses provide nutrients (e.g., carbon [C], nitrogen [N], phosphorus [P]) to freshwater systems (e.g., Juday et al. 1932). More recently, scientists have documented that these “salmon-derived nutrient” subsidies may have significant impacts on both freshwater and riparian communities and on the life histories of organisms that live there (Willson et al. 1998, Cederholm et al. 1999). Because of the burgeoning interest in salmon, growing indications of their ecological importance, and recent calls for management to consider the role of salmon in aquatic and terrestrial ecosystems (e.g., Larkin and Slaney 1997), we take this opportunity to review what is understood about the function of salmon as key elements of ecological systems. Our objectives are twofold. First, we expand on previous reviews of salmon (Willson et al. 1998, Cederholm et al. 1999) to include recent research that has amplified and modified earlier ideas about the contribution of salmon to ecosystem processes. In doing so, we describe the composition, magnitude, and distribution of marine inputs to freshwater and terrestrial systems via salmon. We use an expanding group of studies pertaining to stream nutrient budgets and salmon physiology to construct a schematic that illustrates salmon-derived products and the pathways by which they enter and are retained in aquatic and terrestrial food webs. We then consider the ecological variation associated with salmonid ecosystems and how this may influence the ecological response to the salmon input. Second, we consider how this variation in ecosystem response may influence management and conservation efforts.

Evolutionarily enlightened management

Here we review growing evidence that microevolutionary changes may often be rapid and, in many cases, occur on time frames comparable to human disturbance and anthropogenic change. Contemporary evolutionary change has been documented in relatively pristine habitats, in disturbed populations, under captive management, and in association with both intentional and inadvertent introductions. We argue that evolutionary thinking is thus relevant to conservation biology and resource management but has received insufficient consideration. Ignoring evolution may have a variety of consequences, including unpredicted evolutionary responses to disturbance and naive or inappropriate management decisions. Philosophically, we must also grapple with the issue of whether the evolution of adaptations to disturbance and degraded habitats is sometimes beneficial or something to be rigorously avoided. We advocate promoting evolutionarily enlightened management [Lecture Notes in Biomathematics 99 (1994) 248], in which both the ecological and evolutionary consequences of resource management decisions are considered.

Consumption choice by bears feeding on salmon

Consumption choice by brown (Ursus arctos) and black bears (U. americanus) feeding on salmon was recorded for over 20,000 bear-killed fish from 1994 to 1999 in Bristol Bay (sockeye salmon, Oncorhynchus nerka) and southeastern Alaska (pink, O. gorbuscha and chum salmon O. keta). These data revealed striking patterns of partial and selective consumption that varied with relative availability and attributes of the fish. As the availability of salmon decreased, bears consumed a larger proportion of each fish among both years and habitats. When availability was high (absolute number and density of salmon), bears consumed less biomass per captured fish, targeting energy-rich fish (those that had not spawned) or energy-rich body parts (eggs in females; brain in males). In contrast, individual fish were consumed to a much greater extent, regardless of sex or spawning status, in habitats or years of low salmon availability. The proportion of biomass consumed per fish was similar for males and females, when spawning status was statistically controlled, but bears targeted different body parts: the body flesh, brain and dorsal hump in males and the roe in females. Bears thus appeared to maximize energy intake by modifying the amount and body parts consumed, based on availability and attributes of spawning salmon.

Magnitude and Fate of Salmon-Derived Nutrients and Energy in a Coastal Stream Ecosystem

ABSTRACT We quantified the energy and mineral (nitrogen, phosphorous) composition of live pink salmon (Oncorhynchus gorbuscha) and chum salmon (O. keta), their eggs, and carcasses, and tracked the fate of chum salmon spawning in a small Alaskan coastal stream. On average, salmon entered streams with 5.3 kJ·g-−1, 3.3% N, 0.48% P. Much of the energy in female salmon was stored in the gametes because the gonads were both large (20% of their wet body mass) and high in energy density (11 kJ/g). Carcasses following senescent death had lower mass-specific energy and N (but not P) compared to fish at stream entrance. Bears removed nearly 50% of the salmon-derived nutrients and energy from the stream by capturing salmon and dragging the carcasses from the stream. Much of the salmon biomass was made available to riparian scavengers because bears partially consumed the fish. Nutrients bound in salmon tissue at senescent death were quickly exported to the estuary after only a few days because of periodic high flows and low rates of scavenging by bears.

Transportation of Pacific salmon carcasses from streams to riparian forests by bears

Predation on Pacific salmon by bears (genus Ursus L., 1758) can be an important ecosystem process because the spatial distribution of carcasses largely determines whether marine-derived nutrients cycle through aquatic or terrestrial pathways. Direct observations on three streams in southeastern Alaska indicated that 49% of the pink (Oncorhynchus gorbuscha (Walbaum, 1792)) and chum (Oncorhynchus keta (Walbaum in Artedi, 1792)) salmon killed by bears were carried into the forest. The tendency of bears to transport carcasses was independent of the sex and species of salmon, but unspawned fish were more often transported than fish that had completed spawning. Data on tagged sockeye salmon (Oncorhynchus nerka (Walbaum in Artedi, 1792)) in one southwestern Alaska stream indicated that 42.6% of the killed salmon were transported, and that higher percentages were transported in years when salmon densities were greater. At six other streams, on average, 68.1% of the sockeye salmon killed were apparently transporte...

NESTING SUCCESS OF FOREST BIRDS IN SOUTHEAST ALASKA AND ADJACENT CANADA

Abstract Predation caused 78% of nest failures in coastal forests of southeast Alaska and interior forests of adjacent Canada. Nest success tended to be better in coastal than interior forests. Mayfield daily nest survival from predation on open-cup nests was higher in egg than nestling phase for most species. Species building large (thrush-sized) nests had lower Mayfield daily survival from predation than species building smaller (warbler-sized) nests, but there was no difference in daily survival (total and from predation only) among species nesting in different vegetation strata. Nesting success differed little with nest cover or nest site diversity for most species. Total nest success within species was only sometimes higher in commonly used nest sites than in less frequently used sites. Nest survival from predation did not generally decrease with increasing nest density within guilds of species with similar nests or with nest-site similarity. We emphasize the likelihood of varied outcomes of natural selection on nest-site selection in differing circumstances.

BACKGROUND MATCHING AND COLOR-CHANGE PLASTICITY IN COLONIZING FRESHWATER SCULPIN POPULATIONS FOLLOWING RAPID DEGLACIATION

Anthropogenic-induced change is forcing organisms to shift their distributions and colonize novel habitats at an increasing rate, which leads to complex interactions among evolutionary processes. Coastrange sculpin (Cottus aleuticus) have colonized recently deglaciated streams of Glacier Bay in Alaska within the last 220 years. We examined divergence among populations in background matching coloration and tested the hypothesis that observed variation is due to morphological color plasticity. To examine how color-change plasticity has interacted with other evolutionary processes, we also determined the influence of colonization on neutral genetic diversity. We observed clinal variation in substrate-matching fish color along the chronological continuum of streams. Microsatellites provided little evidence of genetic subdivision among sculpin populations. Fish color was significantly correlated to substrate color, but was not correlated to neutral population genetic structure. Furthermore, a laboratory experiment revealed that morphological color plasticity could explain much, but not all, of the observed fish color divergence. Our study demonstrates that sculpin in Glacier Bay have colonized and adapted to recently deglaciated habitat and suggests that color change plasticity has aided in this process. This research, therefore, highlights the important role phenotypic plasticity may play in the adaptation of species to rapid climate change.

A Bayesian approach for understanding the role of ship speed in whale-ship encounters

Mandatory or voluntary reductions in ship speed are a common management strategy for reducing deleterious encounters between large ships and large whales. This has produced strong resistance from shipping and marine transportation entities, in part because very few studies have empirically demonstrated whether or to what degree ship speed influences ship-whale encounters. Here we present the results of four years of humpback whale sightings made by observers aboard cruise ships in Alaska, representing 380 cruises and 891 ship-whale encounters. Encounters occurred at distances from 21 m to 1000 m (x = 567 m) with 61 encounters (7%) occurring between 200 m and 100 m, and 19 encounters (2%) within 100 m. Encounters were spatially aggregated and highly variable across all ship speeds. Nevertheless a Bayesian change-point model found that the relationship between whale distance and ship speed changed at 11.8 knots (6.1 m/s) with whales encountering ships, on average, 114 m closer when ship speeds were above 11.8 knots. Binning encounter distances by 1-knot speed increments revealed a clear decrease in encounter distance with increasing ship speed over the range of 7-17 knots (3.6-8.7 m/s). Our results are the first to demonstrate that speed influences the encounter distance between large ships and large whales. Assuming that the closer ships come to whales the more likely they are to be struck, our results suggest that reduced ship speed may be an effective management action in reducing the probability of a collision.

PASSERINE DENSITIES IN RIPARIAN FORESTS OF SOUTHEAST ALASKA: POTENTIAL EFFECTS OF ANADROMOUS SPAWNING SALMON

Abstract Spawning salmon (Oncorhynchus spp.) greatly influence many biotic processes in both the aquatic and terrestrial components of riparian systems, possibly including the community of breeding birds. In southeastern Alaska in 1996–1997, the riparian forests bordering salmon streams supported, on average, higher densities, but not diversity, of forest passerines compared to non-salmon streams. The presence of salmon in the fall increases the abundance of terrestrial and aquatic invertebrates and possibly enhances food resources to riparian birds in the spring. Densidades de Aves Paserinas en Bosques Riparios del Sureste de Alaska: Efectos Potenciales del Desove de Salmones Anádromos Resumen. El desove de salmones anádromos (Oncorhynchus spp.) ejerce una importante influencia sobre muchos procesos bióticos en los componentes acuáticos y terrestres de los sistemas ribereños, incluyendo posiblemente las comunidades de aves. Entre 1996 y 1997, los bosques riparios del sureste de Alaska ubicados alrededor de arroyos con salmones tuvieron en promedio mayores densidades (pero no mayor diversidad) de aves paserinas de bosque que bosques situados alrededor de arroyos sin salmones. La presencia de los salmones en el otoño incrementa la abundancia de invertebrados terrestres y acuáticos y posiblemente causa aumentos en la cantidad de recursos alimenticios disponibles para las aves ribereñas en la primavera.

Post-Breeding Season Migrations of a Top Predator, the Harbor Seal (Phoca vitulina richardii), from a Marine Protected Area in Alaska

Marine protected areas (MPAs) are increasingly being used as a conservation tool for highly mobile marine vertebrates and the focus is typically on protecting breeding areas where individuals are aggregated seasonally. Yet movements during the non-breeding season can overlap with threats that may be equally as important to population dynamics. Thus understanding habitat use and movements of species during the non-breeding periods is critical for conservation. Glacier Bay National Park, Alaska, is one of the largest marine mammal protected areas in the world and has the only enforceable protection measures for reducing disturbance to harbor seals in the United States. Yet harbor seals have declined by up to 11.5%/year from 1992 to 2009. We used satellite-linked transmitters that were attached to 37 female harbor seals to quantify the post-breeding season migrations of seals and the amount of time that seals spent inside vs. outside of the MPA of Glacier Bay. Harbor seals traveled extensively beyond the boundaries of the MPA of Glacier Bay during the post-breeding season, encompassing an area (25,325 km2) significantly larger than that used by seals during the breeding season (8,125 km2). These movements included the longest migration yet recorded for a harbor seal (3,411 km) and extended use (up to 23 days) of pelagic areas by some seals. Although the collective utilization distribution of harbor seals during the post-breeding season was quite expansive, there was a substantial degree of individual variability in the percentage of days that seals spent in the MPA. Nevertheless, harbor seals demonstrated a high degree of inter-annual site fidelity (93%) to Glacier Bay the following breeding season. Our results highlight the importance of understanding the threats that seals may interact with outside of the boundaries of the MPA of Glacier Bay for understanding population dynamics of seals in Glacier Bay.