Linking elements of the Integrated Ocean Observing System (IOOS) with the planned National Water Quality Monitoring Network. Proceedings from the NOAA-Supported workshop 19-21 September 2005

The National Oceanic and Atmospheric Administration (NOAA), in cooperation with the New Jersey Marine Sciences Consortium (NJMSC), hosted a workshop at Rutgers University on 19-21 September 2005 to explore ways to link the U.S. Integrated Ocean Observing System (IOOS) to the emerging infrastructure of the National Water Quality Monitoring Network (NWQMN). Participating partners included the Mid-Atlantic Coastal Ocean Observing Regional Association, U.S. Geological Survey, Rutgers University Coastal Ocean Observing Laboratory, and the New Jersey Sea Grant College. The workshop was designed to highlight the importance of ecological and human health linkages in the movement of materials, nutrients, organisms and contaminants along the Delaware Bay watershed-estuary-coastal waters gradient (hereinafter, the “Delaware Bay Ecosystem [DBE]”), and to address specific water quality issues in the mid-Atlantic region, especially the area comprising the Delaware River drainage and near-shore waters. Attendees included federal, state and municipal officials, coastal managers, members of academic and research institutions, and industry representatives. The primary goal of the effort was to identify key management issues and related scientific questions that could be addressed by a comprehensive IOOS-NWQMN infrastructure (US Commission on Ocean Policy 2004; U.S. Ocean Action Plan 2004). At a minimum, cooperative efforts among the three federal agencies (NOAA, USGS and EPA) involved in water quality monitoring were required. Further and recommended by the U.S. Commission on Ocean Policy, outreach to states, regional organizations, and tribes was necessary to develop an efficient system of data gathering, quality assurance and quality control protocols, product development, and information dissemination.

Estuarine use assessment: trial project in Casco Bay, Maine

The National Centers for Coastal Ocean Science (NCCOS) of the National Oceanic and Atmospheric Administration (NOAA) is interested in developing a project to determine the health of estuaries based on the stated or desired uses of society. An estuarine use assessment could complement the National Coastal Assessment, which tracks coastal and estuarine health through a series of environmental indicators. These indicators are used to assign a “score” to each coastal region, with some indicators reflecting the ability of the region to support desired uses such as fishing and swimming. An estuarine use assessment could also provide valuable information to resource managers and other decision-makers as they face decisions about the optimal and most sustainable mix of activities in an estuary. An initial step of an estuarine use assessment would be to define and quantify the desired societal uses of the estuary. Society includes residents living near the estuary or industries relying on the estuary, seasonal residents and tourists that use the estuary on a more limited basis, and the public at-large that may use or value the estuary indirectly. The desired uses may include discrete, visible uses such as swimming, recreational or commercial fishing, and navigation. They also may extend to broader, more intangible uses such as maintaining ecological functions or aesthetic appeal. National legislation such as the Estuary Restoration Act, which promotes and funds the restoration of estuaries in the U.S., reflects the public’s desire for estuaries to retain their ecological structures and functions. This report summarizes a project carried out in 2003 that attempted to quantify the desired human uses of a specific estuary in Maine and to determine current measures of success used by coastal managers in Maine to track the ability of the estuary to support desired uses. Casco Bay was chosen as the spatial embayment for which to delineate uses, and nutrient enrichment was selected as the parameter for confirming assumptions about current measures of outcomes related to uses. The report highlights some of the challenges to completing an estuarine use assessment and offers general recommendations for addressing these challenges.

Protocols for conducting dolphin capture-release health assessment studies

Marine mammals, such as dolphins, can serve as key indicator species in coastal areas by reflecting the effects of natural and anthropogenic stressors. As such they are often considered sentinels of environmental and ecosystem health (Bossart 2006; Wells et al. 2004; Fair and Becker 2000). The bottlenose dolphin is an apex predator and a key component of many estuarine environments in the southeastern United States (Woodward-Clyde Consultants 1994; SCDNR 2005). Health assessments of dolphins are especially critical in areas where populations are depleted, show signs of epidemic disease and/or high mortality and/or where habitat is being altered or impacted by human activities. Recent assessments of environmental conditions in the Indian River Lagoon, Florida (IRL) and the estuarine waters surrounding Charleston, South Carolina (CHS) highlight the need for studies of the health of local bottlenose dolphins. While the condition of southeastern estuaries was rated as fair in the National Coastal Condition Report (U.S. EPA 2001), it was noted that the IRL was characterized by poorer than expected benthic communities, significant sediment toxicity and increased nutrient concentrations. Similarly, portions of the CHS estuary have sediment concentrations of aliphatic aromatic hydrocarbons, select inorganic metals, and some persistent pesticides far in excess of reported bioeffect levels (Hyland et al. 1998). Long-term trends in water quality monitoring and recent scientific research suggest that waste load assimilation, non-point source runoff impacts, contaminated sediments, and toxic pollutants are key issues in the CHS estuary system. Several ‘hot spots’ with high levels of heavy metals and organic compounds have been identified (Van Dolah et al. 2004). High concentrations of anthropogenic trace metals, polychlorinated biphenyls (PCB’s) and pesticides have been found in the sediments of Charleston Harbor, as well as the Ashley and Cooper Rivers (Long et al. 1998). Two superfund sites are located within the CHS estuary and the key contaminants of concern associated with these sites are: polycyclic aromatic hydrocarbons (PAH), lead, chromium, copper, arsenic, zinc and dioxin. Concerns related to the overall health of IRL dolphins and dermatologic disease observed in many dolphins in the area (Bossart et al. 2003) initiated an investigation of potential factors which may have impacted dolphin health. From May-August 2001, 35 bottlenose dolphins died in the IRL during an unusual mortality event (MMC 2003). Many of these dolphins were diagnosed with a variety of skin lesions including proliferative ulcerative dermatitis due to protozoa and fungi, dolphin pox and a vesicular dermatopathy of unknown etiology (Bossart et al. 2003). Multiple species from fish to dolphins in the IRL system have exhibited skin lesions of various known and unknown etiologies (Kane et al. 2000; Bossart et al. 2003; Reif et al. 2006). On-going photo-identification (photo-ID) studies have documented skin diseases in IRL dolphins (Mazzoil et al. 2005). In addition, up to 70% of green sea turtles in the IRL exhibit fibropapillomas, with the highest rates of occurrence being seen in turtles from the southern IRL (Hirama 2001).

The effects of urbanization on populations of grass shrimp Palaemonetes spp. in small, high salinity estuaries

High salinity estuaries in the southeastern U.S. have experienced increased inputs of contaminants from nonpoint source (NPS) urban runoff and decreases in habitat due to filling of wetlands and dock/bulkhead construction. Urbanization may pose significant risks to estuarine fauna, particularly crustaceans. The grass shrimp of the genus Palaemonetes, is one of the dominant species found in estuarine tidal creeks, accounting for greater than 50% of all macropelagic fauna on an annual basis. Spatial analytical and geographic information system techniques were used to determine which factors influenced the Palaemonetes population structures in a South Carolina bar-built estuary surrounded by urban development. Impacts from land use practices were investigated using concentric circular buffers around study sites. Factors investigated included sediment-associated polycyclic aromatic hydrocarbons concentration, land use classification, percent impervious surfaces, and other selected urban factors. Geographic information system and statistical modeling showed quantitative relationships between land use class and impacts on Palaemonetes density. The study suggests that habitat loss is a major factor influencing grass shrimp densities. Multiple regression modeling suggests a significant relationship between habitat alterations and Palaemonetes densities.

Baseline assessment of Guánica Bay, Puerto Rico in support of watershed restoration

Guánica Bay is a major estuary on the southwest coast of Puerto Rico. Significant coral reef ecosystems are present outside the bay. These valuable habitats may be impacted by transport of sediments, nutrients and contaminants from the watershed, through the bay and into the offshore waters. The National Oceanic and Atmospheric Administration’s (NOAA) National Centers for Coastal Ocean Science (NCCOS), in consultation with local and regional experts, conducted an interdisciplinary assessment of coral reef ecosystems, contaminants, sedimentation rates and nutrient distribution patterns in and around Guánica Bay. This work was conducted using many of the same protocols as ongoing monitoring work underway elsewhere in the U.S. Caribbean and has enabled comparisons among coral reef ecosystems between this study and other locations in the region. This characterization of Guánica marine ecosystems establishes benchmark conditions that can be used for comparative documentation of future change, including possible negative outcomes due to future land use change, or improvement in environmental conditions arising from management actions. This report is organized into six chapters that represent a suite of interrelated studies. Chapter 1 provides a short introduction to the study area. Chapter 2 is focused on biogeographic assessments and benthic mapping of the study area, including new surveys of fish, marine debris and reef communities on hardbottom habitats in the study area. Chapter 3 quantifies the distribution and magnitude of a suite of contaminants (e.g., heavy metals, PAHs, PCBs, pesticides) in both surface sediments and coral tissues. Chapter 4 presents results of sedimentation measurements in and outside of the bay. Chapter 5 examines the distribution of nutrients in in the bay, offshore from the bay and in the watershed. Chapter 6 is a brief summary discussion that highlights key findings of the entire suite of studies.

National Centers for Coastal Ocean Science (NCCOS) research highlights in the Chesapeake Bay

The Chesapeake Bay is the largest estuary in the United States. It is a unique and valuable national treasure because of its ecological, recreational, economic and cultural benefits. The problems facing the Bay are well known and extensively documented, and are largely related to human uses of the watershed and resources within the Bay. Over the past several decades as the origins of the Chesapeake’s problems became clear, citizens groups and Federal, State, and local governments have entered into agreements and worked together to restore the Bay’s productivity and ecological health. In May 2010, President Barack Obama signed Executive Order number 13508 that tasked a team of Federal agencies to develop a way forward in the protection and restoration of the Chesapeake watershed. Success of both State and Federal efforts will depend on having relevant, sound information regarding the ecology and function of the system as the basis of management and decision making. In response to the executive order, the National Oceanic and Atmospheric Administration’s National Centers for Coastal Ocean Science (NCCOS) has compiled an overview of its research in Chesapeake Bay watershed. NCCOS has a long history of Chesapeake Bay research, investigating the causes and consequences of changes throughout the watershed’s ecosystems. This document presents a cross section of research results that have advanced the understanding of the structure and function of the Chesapeake and enabled the accurate and timely prediction of events with the potential to impact both human communities and ecosystems. There are three main focus areas: changes in land use patterns in the watershed and the related impacts on contaminant and pathogen distribution and concentrations; nutrient inputs and algal bloom events; and habitat use and life history patterns of species in the watershed. Land use changes in the Chesapeake Bay watershed have dramatically changed how the system functions. A comparison of several subsystems within the Bay drainages has shown that water quality is directly related to land use and how the land use affects ecosystem health of the rivers and streams that enter the Chesapeake Bay. Across the Chesapeake as a whole, the rivers that drain developed areas, such as the Potomac and James rivers, tend to have much more highly contaminated sediments than does the mainstem of the Bay itself. In addition to what might be considered traditional contaminants, such as hydrocarbons, new contaminants are appearing in measurable amounts. At fourteen sites studied in the Bay, thirteen different pharmaceuticals were detected. The impact of pharmaceuticals on organisms and the people who eat them is still unknown. The effects of water borne infections on people and marine life are known, however, and the exposure to certain bacteria is a significant health risk. A model is now available that predicts the likelihood of occurrence of a strain of bacteria known as Vibrio vulnificus throughout Bay waters.

Severe droughts reduce estuarine primary productivity with cascading effects on higher trophic levels

Using a 10-yr time-series data set, we analyzed the effects of two severe droughts on water-quality and ecosystem processes in a temperate, eutrophic estuary (Neuse River Estuary, North Carolina). During the droughts, dissolved inorganic nitrogen concentrations were on average 46–68% lower than the long-term mean due to reduced riverine input. Phytoplankton productivity and biomass were slightly below average for most of the estuary during a spring–autumn drought in 2002, but were dramatically lower than average throughout the estuary during an autumn–winter drought in 2007–2008. Droughts affected upper trophic levels through alteration of both habitat condition (i.e., bottom-water dissolved oxygen levels) and food availability. Bottomwater dissolved oxygen levels were near or slightly above average during the 2002 drought and during summer 2007. Concomitant with these modest improvements in bottom-water oxygen condition, fish kills were greatly reduced relative to the long-term average. Low-oxygen bottom-water conditions were more pronounced during summer 2008 in the latter stages of the 2007–2008 drought, and mesozooplankton abundances were eight-fold lower in summer 2008 than during nondrought years. Below-average mesozooplankton abundances persisted for well over 1 yr beyond cessation of the drought. Significant fish kills were observed in summer 2008 and 2009, perhaps due to the synergistic effects of hypoxia and reduced food availability. These results indicate that droughts can exert both ephemeral and prolonged multiyear influence on estuarine ecosystem processes and provide a glimpse into the future, when many regions of the world are predicted to face increased drought frequency and severity due to climate change.

Weighing your options: how to protect your property from shoreline erosion, a handbook for estuarine property owners in North Carolina

If you own property on one of North Carolina’s estuaries, you can use this guide as a tool to learn about the choices you have to control your shoreline erosion and help decide which approach may be right for you. In North Carolina, we make a distinction between waterfront property that is located on the estuary, referred to as estuarine, shoreline, soundfront or riverside property, and waterfront property located directly on the ocean, referred to as oceanfront. Why? State laws and regulations addressing estuarine and oceanfront property, and the available erosion control methods, are quite different. This guide focuses on estuarine property. We’ll introduce you to the six main erosion control options in use in North Carolina and give you information about the out-of-pocket costs and tangible benefits of each option. We’ll also give you information about “hidden” costs and benefits that you may want to factor into your decision-making. You are fortunate to have a piece of estuarine shoreline to call your own, whether it’s your year-round residence or a weekend getaway. And if you’ve noticed some shoreline erosion lately, you’re probably a little concerned. But there are ready solutions.

Using wildlife activity and antibiotic resistance analysis to model bacterial water quality in coastal ponds

Models that help predict fecal coliform bacteria (FCB) levels in environmental waters can be important tools for resource managers. In this study, we used animal activity along with antibiotic resistance analysis (ARA), land cover, and other variables to build models that predict bacteria levels in coastal ponds that discharge into an estuary. Photographic wildlife monitoring was used to estimate terrestrial and aquatic wildlife activity prior to sampling. Increased duck activity was an important predictor of increased FCB in coastal ponds. Terrestrial animals like deer and raccoon, although abundant, were not significant in our model. Various land cover types, rainfall, tide, solar irradiation, air temperature, and season parameters, in combination with duck activity, were significant predictors of increased FCB. It appears that tidal ponds allow for settling of bacteria under most conditions. We propose that these models can be used to test different development styles and wildlife management techniques to reduce bacterial loading into downstream shellfish harvesting and contact recreation areas.

Historical nitrogen and phosphorus loadings to the northern Gulf of Mexico

Primary productivity in many coastal systems is nitrogen (N) limited; although, phytoplankton productivity may be limited by phosphorus (P) seasonally or in portions of an estuary. Increases in loading of limiting nutrients to coastal ecosystems may lead to eutrophication (Nixon 1996). Anthropogenically enhanced eutrophication includes symptoms such as loss of seagrass beds, changes in algal community composition, increased algal (phytoplankton) blooms (Richardson et al. 2001), hypoxic or anoxic events, and fish kills (Bricker et al. 2003).

Characterizing Jobos Bay, Puerto Rico: a watershed modeling analysis and monitoring plan

Land-based pollution is commonly identified as a major contributor to the observed deterioration of shallow-water coral reef ecosystem health. Human activity on the coastal landscape often induces nutrient enrichment, hypoxia, harmful algal blooms, toxic contamination and other stressors that have degraded the quality of coastal waters. Coral reef ecosystems throughout Puerto Rico, including Jobos Bay, are under threat from coastal land uses such as urban development, industry and agriculture. The objectives of this report were two-fold: 1. To identify potentially harmful land use activities to the benthic habitats of Jobos Bay, and 2. To describe a monitoring plan for Jobos Bay designed to assess the impacts of conservation practices implemented on the watershed. This characterization is a component of the partnership between the U.S. Department of Agriculture (USDA) and the National Oceanic and Atmospheric Administration (NOAA) established by the Conservation Effects Assessment Project (CEAP) in Jobos Bay. CEAP is a multi-agency effort to quantify the environmental benefits of conservation practices used by private landowners participating in USDA programs. The Jobos Bay watershed, located in southeastern Puerto Rico, was selected as the first tropical CEAP Special Emphasis Watershed (SEW). Both USDA and NOAA use their respective expertise in terrestrial and marine environments to model and monitor Jobos Bay resources. This report documents NOAA activities conducted in the first year of the three-year CEAP effort in Jobos Bay. Chapter 1 provides a brief overview of the project and background information on Jobos Bay and its watershed. Chapter 2 implements NOAA’s Summit to Sea approach to summarize the existing resource conditions on the watershed and in the estuary. Summit to Sea uses a GIS-based procedure that links patterns of land use in coastal watersheds to sediment and pollutant loading predictions at the interface between terrestrial and marine environments. The outcome of Summit to Sea analysis is an inventory of coastal land use and predicted pollution threats, consisting of spatial data and descriptive statistics, which allows for better management of coral reef ecosystems. Chapters 3 and 4 describe the monitoring plan to assess the ecological response to conservation practices established by USDA on the watershed. Jobos Bay is the second largest estuary in Puerto Rico, but has more than three times the shoreline of any other estuarine area on the island. It is a natural harbor protected from offshore wind and waves by a series of mangrove islands and the Punta Pozuelo peninsula. The Jobos Bay marine ecosystem includes 48 km² of mangrove, seagrass, coral reef and other habitat types that span both intertidal and subtidal areas. Mapping of Jobos Bay revealed 10 different benthic habitats of varying prevalence, and a large area of unknown bottom type covering 38% of the entire bay. Of the known benthic habitats, submerged aquatic vegetation, primarily seagrass, is the most common bottom type, covering slightly less than 30% of the bay. Mangroves are the dominant shoreline feature, while coral reefs comprise only 4% of the total benthic habitat. However, coral reefs are some of the most productive habitats found in Jobos Bay, and provide important habitat and nursery grounds for fish and invertebrates of commercial and recreational value.

Examination of the foraging habits of Pacific harbor seal (Phoca vitulina richardsi) to describe their use of the Umpqua River, Oregon, and their predation on salmonids

The increase in harbor seal (Phoca vitulina richardsi) abundance, concurrent with the decrease in salmonid (Oncorhynchus spp.) and other fish stocks, raises concerns about the potential negative impact of seals on fish populations. Although harbor seals are found in rivers and estuaries, their presence is not necessarily indicative of exclusive or predominant feeding in these systems. We examined the diet of harbor seals in the Umpqua River, Oregon, during 1997 and 1998 to indirectly assess whether or not they were feeding in the river. Fish otoliths and other skeletal structures were recovered from 651 scats and used to identify seal prey. The use of all diagnostic prey structures, rather than just otoliths, increased our estimates of the number of taxa, the minimum number of individuals and percent frequency of occurrence (%FO) of prey consumed. The %FO indicated that the most common prey were pleuronectids, Pacific hake (Merluccius productus), Pacific stag-horn sculpin (Leptocottus armatus), osmerids, and shiner surfperch (Cymatogaster aggregata). The majority (76%) of prey were fish that inhabit marine waters exclusively and fish found in marine and estuarine areas (e.g. anadromous spp.) which would indicate that seals forage predominantly at sea and use the estuary for resting and opportunistic feeding. Salmonid remains were encountered in 39 samples (6%); two samples contained identifiable otoliths, which were determined to be from chi-nook salmon (O. tshawytscha). Because of the complex salmonid composition in the Umpqua River, we used molecular genetic techniques on salmonid bones retrieved from scat to discern species that were rare from those that were abundant. Of the 37 scats with salmonid bones but no otoliths, bones were identified genetically as chinook or coho (O. kisutch) salmon, or steelhead trout (O. mykiss) in 90% of the samples.

Use of ocean and estuarine habitats by young-of-year bluefish (Pomatomus saltatrix) in the New York Bight

Young-of-year (YOY) blue-fish (Pomatomus saltatrix) along the U.S. east coast are often assumed to use estuaries almost exclusively during the summer. Here we present data from 1995 to 1998 indicating that YOY (30–260 mm FL) also use ocean habitats along the coast of New Jersey. An analysis of historical and recent data on northern and southern ocean beaches (0.1–2 m) and the inner continental shelf (5–27 m) during extensive sampling in New Jersey waters from 1995 to 1998 indicated that multiple cohorts occurred (June–August) in every year. When comparable collections of YOY were made in the ocean and in an adjacent estuary, the abundance was 1–2 orders of magnitude greater on ocean beaches during the summer. The YOY were even more abundant in ocean habitats in the fall (September–October), presumably as a result of YOY leaving estuaries to join the coastal migration south. During 1999 and 2000, YOY bluefish were tagged with internal sequential coded wire microtags in order to refine our under-standing of habitat use and movement. Few (0.04%) of the fish tagged on ocean beaches were recaptured; however, 2.2% of the fish tagged in the estuary were recaptured from 2 to 27 days after tagging. Recaptured fish grew quickly (average 1.37 mm FL/d). On ocean beaches YOY fed on a variety of invertebrates and fishes but their diet changed with size. By approximately 80–100 mm FL, they were piscivorous and fed primarily on engraulids, a pattern similar to that reported in estuaries. Based on distribution, abundance, and feeding, both spring- and summer-spawned cohorts of YOY bluefish commonly use ocean habitats. Therefore, attempts to determine factors affecting recruitment success based solely on estuarine sampling may be inadequate and further examination, especially of the contribution of the summer-spawned cohort in ocean habitats, appears warranted.

Growth, recruitment, and abundance of juvenile striped mullet (Mugil cephalus) in South Carolina estuaries

Growth, recruitment, and abundance of young-of-the-year (YOY) striped mullet (Mugil cephalus L.) in estuarine habitats in South Carolina from 1998 to 2000 were examined and compared to historical data (1986–91) of growth, recruitment, and abundance. Daily growth increments from the sagittal otoliths of juvenile striped mullet were validated by using fish immersed in oxytetracycline hydrochloride (OTC) for five hours from the Charleston Harbor Estuary system. The distribution of back-calculated birthdates indicated that striped mullet spawn from October to late April and estuarine recruitment occurs from January through May. Juveniles were more abundant in mesohaline and polyhaline salinity regimes but were found throughout the estuary. Juvenile growth after recruitment into the estuary can be described by the relationship Total length (mm) = 0.341 (Age)1.04 (r2=0.741, P=0.001). Growth of juveniles according to the analysis of size-frequency data from historical surveys (1986 to 1991) in the same estuaries gave the relationship Total length (mm) = 8.77 (month)1.12 (r2=0.950, P=0.001). The similarity in the growth curves for both groups of fish suggests that juvenile striped mullet in South Carolina have consistent annual growth during the first year of life.

The relative value of different estuarine nursery areas in North Carolina for transient juvenile marine fishes

Offshore winter-spawned fishes dominate the nekton of south-eastern United States estuaries. Their juveniles reside for several months in shallow, soft bottom estuarine creeks and bays called primary nursery areas. Despite similarity in many nursery characteristics, there is, between and within species, variability in the occupation of these habitats. Whether all occupied habitats are equally valuable to individuals of the same species or whether most recruiting juveniles end up in the best habitats is not known. If nursery quality varies, then factors controlling variation in pre-settlement fish distribution are important to year-class success. If nursery areas have similar values, interannual variation in distribution across nursery creeks should have less effect on population sizes or production. I used early nursery period age-specific growth and mortality rates of spot (Leiostomus xanthurus) and Atlantic croaker (Micropogonias undulatus)—two dominant estuarine fishes—to assess relative habitat quality across a wide variety of nursery conditions, assuming that fish growth and mortality rates were direct reflections of overall physical and biological conditions in the nurseries. I tested the hypothesis that habitat quality varies for these fishes by comparing growth and mortality rates and distribution patterns across a wide range of typical nursery habitats at extreme ends of two systems. Juvenile spot and Atlantic croaker were collected from 10 creeks in the Cape Fear River estuary and from 18 creeks in the Pamlico Sound system, North Carolina, during the 1987 recruitment season (mid-March–mid-June). Sampled creeks were similar in size, depth, and substrates but varied in salinities, tidal regimes, and distances from inlets. Spot was widely distributed among all the estuarine creeks, but was least abundant in the creeks in middle reaches of both systems. Atlantic croaker occurred in the greatest abundance in oligohaline creeks of both systems. Instantaneous growth rates derived from daily otolith ages were generally similar for all creeks and for both species, except that spot exhibited a short-term growth depression in the upriver Pamlico system creeks—perhaps the result of the long migration distance of this species to this area. Spot and Atlantic croaker from upriver oligohaline creeks exhibited lower mortality rates than fish from downstream polyhaline creeks. These results indicated that even though growth was similar at the ends of the estuaries, the upstream habitats provided conditions that may optimize fitness through improved survival.