Research

Publications

Below is a list of publications focused on river herring, including their cultural and social importance, life history and population ecology, coastal food webs, threats, and monitoring methods.

These publications can also be found in this public Zotero library.

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Synthesis Papers

Hare, J. A., Borggaard, D. L., Alexander, M. A., Bailey, M. M., Bowden, A. A., Damon-Randall, K., Didden, J. T., Hasselman, D. J., Kerns, T., McCrary, R., McDermott, S., Nye, J. A., Pierce, J., Schultz, E. T., Scott, J. D., Starks, C., Sullivan, K., & Beth Tooley, M. (2021). A Review of River Herring Science in Support of Species Conservation and Ecosystem Restoration. 

Synthesizes current knowledge of river herring, with an emphasis on identification of threats, discussion of recent research and management actions, and identification of research needs.

Hayden, A., Steinman, M., & Gorich, R. (2019). Up and up: River Herring in Eastern Maine. Downeast Fisheries Partnership.

An update on the status of river herring runs in eastern Maine, including the value of river herring and some of the many projects underway to restore access to their historic spawning habitat.

Kritzer, J. P., Hall, C. J., Hoppe, B., Ogden, C., & Cournane, J. M. (2022). Managing Small Fish at Large Scales: The Emergence of Regional Policies for River Herring in the Eastern United States.

Retrospective overview of the emergence of large-scale policy frameworks for river herring across their range in U.S. waters.

Maine Indian Tribal State Commission. (2022). Sea Run: A Study Regarding the Impact of Maine Policies on the Quality and Quantity of Traditional Tribal Fish Stocks and Sustenance Practices.

Ouellet, V., Collins, M. J., Kocik, J. F., Saunders, R., Sheehan, T. F., Ogburn, M. B., & Trinko Lake, T. (2022). The diadromous watersheds-ocean continuum: Managing diadromous fish as a community for ecosystem resilience.

Literature synthesis of ecosystem services provided by diadromous fish and of the current knowledge on their socioeconomic and ecological importance. Highlights critical research gaps impeding current management efforts. Framework to advance holistic management of diadromous species and habitats across the watersheds-ocean continuum to strengthen efforts to restore the productivity of these ecosystems and the services they provide.

Social and Cultural Importance of River Herring

Bassett, E. (2015). Cultural Importance of River Herring to the Passamaquoddy People. Sipayik Environmental Department. 

For over twelve thousand years River Herring (Alewife and Blueback Herring) have migrated from the sea by the millions to spawn in the upper headwaters of the St. Croix River Watershed. Without this fish the Passamaquoddy People would not have survived. This paper is intended to explain the cultural importance of river herring from the Passamaquoddy perspective. 

McClenachan, L., S. Lovell, and C. Keaveney. (2015). Social benefits of restoring historical ecosystems and fisheries: alewives in Maine.

There are many social benefits conferred by the restoration of fish passage and small-scale fisheries, and a positive feedback loop between alewife harvest, community engagement, and motivation for further restoration.

Life History and Population Ecology

Dalton, R. M., Sheppard, J. J., Finn, J. T., Jordaan, A., & Staudinger, M. D. (2022). Phenological Variation in Spring Migration Timing of Adult Alewife in Coastal Massachusetts.

Winter sea surface temperature, spring and fall transition dates, and annual run size were the strongest predictors of river herring run initiation and median dates, while a combination of within-season and seasonal-lag effects influenced run end and duration timing.

Davis, J. P., & Schultz, E. T. (2006). Assessment of Anadromous Alewife and Blueback Herring Populations in Connecticut Coastal Streams and Connecticut River Tributaries. 

River herring population data from two rivers was compared with data collected 40 years ago and found an overall decrease in abundance, greater proportions of younger fish (< 5 years old) and first-time spawners, fewer age classes, and decreased abundance of older fish (> 5 years old).

Davis, J. P., & Schultz, E. T. (2009). Temporal Shifts in Demography and Life History of an Anadromous Alewife Population in Connecticut. 

Recent alewife runs at Bride Brook featured lower abundance and younger, smaller fish, indicating a shift in life history. The results of this study suggest recent increases in predatory pressure or bycatch mortality as promising hypotheses that merit further investigation.

Durbin, A. G., Nixon, S. W., & Oviatt, C. A. (1979). Effects of the Spawning Migration of the Alewife, Alosa Pseudoharengus, on Freshwater Ecosystems.

Alewives contribute significant nutrient inputs to freshwater systems, increasing primary production and supporting insect and benthic invertebrates. 

Gahagan, B. I., Gherard, K. E., & Schultz, E. T. (2010). Environmental and Endogenous Factors Influencing Emigration in Juvenile Anadromous Alewives. 

Pulses of juvenile alewife migration were associated with precipitation events, decreases in water temperature, and increases in stream discharge. Migrating juveniles were, on average, larger, older, faster-growing, and in better condition than nonmigrating juveniles. 

Kellogg, R. L. (1982). Temperature Requirements for the Survival and Early Development of the Anadromous Alewife. 

A study of temperature effects on alewife eggs found that the maximum hatching success occurred at 20.8°C and increased temperatures shortened the average time to hatch (7.4 days at 12.7°C to 3 days at 23.8°C). After hatching, the higher the temperature the more daily larva weight gained and the young alewives preferred 26.3°C in a thermal gradient.

Legett, H. D., Jordaan, A., Roy, A. H., Sheppard, J. J., Somos‐Valenzuela, M., & Staudinger, M. D. (2021). Daily patterns of river herring (Alosa spp.) spawning migrations: environmental drivers and variation among coastal streams in Massachusetts.

Long-term fish counts (8–28 years) from 12 coastal Massachusetts streams indicate that daily change in water temperature is the most consistent predictor of both daily river herring presence–absence and abundance during migration.

McBride, M. C., Hasselman, D. J., Willis, T. V., Palkovacs, E. P., & Bentzen, P. (2015). Influence of stocking history on the population genetic structure of anadromous alewife (Alosa pseudoharengus) in Maine rivers. 

Alewives were found to present locally distinct genetic structures, indicating that stock transfers pose risk for alewife populations to adapt, persist, and evolve due to the decline in genetic differentiation among stock populations.

McCartin, K., Jordaan, A., Sclafani, M., Cerrato, R., & Frisk, M. G. (2019). A New Paradigm in Alewife Migration: Oscillations between Spawning Grounds and Estuarine Habitats. 

Deviations from the typical single upstream migration pattern were found in alewives, such as multiple runs back and forth from estuarine habitats to spawning grounds.

Nelson, G. A., Gahagan, B. I., Armstrong, M. P., Jordaan, A., & Bowden, A. (2020). A life cycle simulation model for exploring causes of population change in Alewife (Alosa pseudoharengus). 

A first of its kind full life-cycle model for alewives can be used to understand how external factors and management actions can affect the growth, survival, and migration of alewives.

Ogburn, M. B., Hasselman, D. J., Schultz, T. F., & Palkovacs, E. P. (2017). Genetics and Juvenile Abundance Dynamics Show Congruent Patterns of Population Structure for Depleted River Herring Populations in the Upper Chesapeake Bay. 

Genetic analysis of river herring in the Chesapeake Bay found at least two genetically distinguishable groups of spawning populations for alewives, and at least three for blueback herring, which should be considered separately for conservation and management. 

Palkovacs, E. P., Hasselman, D. J., Argo, E. E., Gephard, S. R., Limburg, K. E., Post, D. M., Schultz, T. F., & Willis, T. V. (2014). Combining genetic and demographic information to prioritize conservation efforts for anadromous alewife and blueback herring. 

Using population genetic data and demographic information, three distinct stocks of alewife and four stocks of blueback herring were identified along the Atlantic coast. The most severe population declines have occurred for populations in the Southern New England and Mid-Atlantic Stocks. 

Reid, K., Carlos Garza, J., Gephard, S. R., Caccone, A., Post, D. M., & Palkovacs, E. P. (2020). Restoration-mediated secondary contact leads to introgression of alewife ecotypes separated by a colonial-era dam. 

After the removal of a colonial-era dam, the reintroduction of anadromous alewives with landlocked alewives resulted in the directional introgression (gene flow between previously diverged lineages) of anadromous alleles in the landlocked population. This reintroduction effort also found that anadromous and landlocked alewife distribute differently in littoral and pelagic habitats.

Reid, K., Palkovacs, E. P., Hasselman, D. J., Baetscher, D., Kibele, J., Gahagan, B., Bentzen, P., McBride, M. C., & Garza, J. C. (2018). Comprehensive evaluation of genetic population structure for anadromous river herring with single nucleotide polymorphism data.

Single nucleotide polymorphism data was collected from 8000 river herring along their entire Atlantic coast range in order to develop a database that can assign stock individuals to their regional genetic group of origin.

Rillahan, C. and He, P. (2023). Waiting for the right time and tide: The fine-scale migratory behavior of river herring in two coastal New England streams.

The changing relationship between the time of day and tidal state within the season manifested in changing periodicity in fish movement to correlate with favorable movement conditions. Sampling methodologies that collect information during all 24 h would likely produce the most accurate run size estimates.

Rosset, J., Roy, A. H., Gahagan, B. I., Whiteley, A. R., Armstrong, M. P., Sheppard, J. J., & Jordaan, A. (2017). Temporal Patterns of Migration and Spawning of River Herring in Coastal Massachusetts. 

A comparison of estimated spawn dates with migration runs found a delay between the start of migration the spawning period within 20 coastal Massachusetts lakes. This supports recent studies that suggest alewives are indeterminate spawners (meaning eggs can develop at any time during the spawning season.)

Stevens, J. R., Saunders, R., & Duffy, W. (2021). Evidence of Life Cycle Diversity of River Herring in the Penobscot River Estuary, Maine. 

The use of the Penobscot River estuary during April through September by one to two year old river herring fish furthers the building evidence that the alewife and blueback herring exhibit high life cycle diversity and “non textbook” migration patterns.

Turner, S. M., & Limburg, K. E. (2016). Juvenile river herring habitat use and marine emigration trends: Comparing populations. 

The chemical analysis of otoliths combined with otolith growth models found small differences in juvenile nursery habitat selection between alewives and blueback herrings. Estuarine nursery use was also found to be more common at lower latitudes.

Turner, S. M., Limburg, K. E., & Palkovacs, E. P. (2015). Can different combinations of natural tags identify river herring natal origin at different levels of stock structure?

Combining otolith chemistry and genetics can help trace river herring in marine bycatch back to its river of origin.

Webb, A. E. (2021). Juvenile alewife (Alosa pseudoharengus) feeding habits, movement and residency in a northern temperate estuary.

The Penobscot Estuary serves as a significant feeding habitat for juvenile alewife because of the presence of calanoid copepods, barnacle larvae, and mysid shrimp.

Wynne, M. L. P., Wilson, K. A., & Limburg, K. E. (2015). Retrospective examination of habitat use by blueback herring (Alosa aestivalis) using otolith microchemical methods. 

Blueback herring otolith microchemistry and concentrations of metals found in water were used to determine patterns in habitat use. The majority exhibited a longer stay in freshwater or low salinity habitats compared to some fish who migrated into seawater in under a year.

Coastal Food Webs and Interactions

Bethoney, N. D., Stokesbury, K. D. E., Schondelmeier, B. P., Hoffman, W. S., & Armstrong, M. P. (2014). Characterization of River Herring Bycatch in the Northwest Atlantic Midwater Trawl Fisheries. 

Bycatch assessment from New Jersey to Southern Maine found that in the northern areas mostly mature river herring were caught, and in the southern areas, mostly juvenile and pre-spawning adults, and migratory adults were caught.

Cournane, J. M., Kritzer, J. P., & Correia, S. J. (2013). Spatial and temporal patterns of anadromous alosine bycatch in the US Atlantic herring fishery. 

Bycatch in the Atlantic herring fishery is more likely to occur in the winter in Southern New England to Mid Atlantic Bight waters, and in the spring, bycatch can occur from the Gulf of Maine to the Mid-Atlantic Bight.

Dias, B. S., Frisk, M. G., & Jordaan, A. (2019). Opening the tap: Increased riverine connectivity strengthens marine food web pathways. 

This study highlights the benefits of increased connectivity between freshwater and ocean ecosystems, including the significant role that anadromous forage fish could play in improving specific fisheries, including Atlantic cod. 

Dias, B. S., Frisk, M. G., & Jordaan, A. (2021). Contrasting fishing effort reduction and habitat connectivity as management strategies to promote alewife (Alosa pseudoharengus) recovery using an ecosystem model. 

Recovery of anadromous forage fish through increased connectivity would build ecosystem resilience to climate, fisheries, and other perturbations. 

Hasselman, D. J., Anderson, E. C., Argo, E. E., Bethoney, N. D., Gephard, S. R., Post, D. M., Schondelmeier, B. P., Schultz, T. F., Willis, T. V., & Palkovacs, E. P. (2015). Genetic stock composition of marine bycatch reveals disproportional impacts on depleted river herring genetic stocks. 

Bycatch of alewives and blueback herring in the southern New England Atlantic herring fishery may be negatively impacting river herring recovery efforts. 

McDermott, S. P., Bransome, N. C., Sutton, S. E., Smith, B. E., Link, J. S., & Miller, T. J. (2015). Quantifying alosine prey in the diets of marine piscivores in the Gulf of Maine: Alosine prey in marine piscivores. 

Alewife, blueback herring, and American shad were found in the diets of common marine piscivores in the northwest Atlantic Ocean, with prey consumption concentrated in near-coastal waters.

Scopel, L. C., Diamond, A. W., Kress, S. W., Hards, A. R., & Shannon, P. (2018). Seabird diets as bioindicators of Atlantic herring recruitment and stock size: A new tool for ecosystem-based fisheries management. 

Seabird diets were used to suggest spatial structuring and movement patterns of Atlantic herring before recruitment into the fishery.

Smith, K. M., Byron, C. J., & Sulikowski, J. A. (2016). Modeling Predator–Prey Linkages of Diadromous Fishes in an Estuarine Food Web. 

A food web model for the Saco River estuary determined the trophic roles of fish; the model suggested that juvenile marine transients (e.g., non-diadromous fish species) are more likely to be foraged than the alewives and blueback herring.

Turner, S. M., Hare, J. A., Richardson, D. E., & Manderson, J. P. (2017). Trends and Potential Drivers of Distribution Overlap of River Herring and Commercially Exploited Pelagic Marine Fishes on the Northeast U.S. Continental Shelf. 

Incidental catches of river herring in commercial Atlantic herring and Atlantic mackerel fisheries are due to the expansion of these species’ distribution caused by increasing abundance. The overlap of co-occurring species’ changes by species’ response rate to temperature changes.

Willis, T. V., Wilson, K. A., Alexander, K. E., & Leavenworth, W. B. (2013). Tracking cod diet preference over a century in the northern Gulf of Maine: Historic data and modern analysis.

Atlantic cod diets from the early 2000’s resemble diets from the late 1890’s which consisted of mainly invertebrates due to the lack of forage fish. Adult cod diets in 195 were primarily composed of fish (mostly Atlantic herring) during a period when dredge and trawl gear usage impacted the benthos in Passamaquoddy Bay.  

Threats

Lynch, P. D., Nye, J. A., Hare, J. A., Stock, C. A., Alexander, M. A., Scott, J. D., Curti, K. L., & Drew, K. (2015). Projected ocean warming creates a conservation challenge for river herring populations. 

Climate change is expected to reduce suitable habitat for river herring across the US Atlantic coast, and alter the marine distribution of both alewives and blueback herring.

Zydlewski, J., Stich, D. S., Roy, S., Bailey, M., Sheehan, T., & Sprankle, K. (2021). What Have We Lost? Modeling Dam Impacts on American Shad Populations Through Their Native Range. 

Through modeling, it is estimated that the American shad has experienced a 41% loss in habitat connectivity because of the damming of coastal systems across the species’ range.

Monitoring, Citizen Science. and Sampling Methods

Bieluch, K. H., Willis, T., Smith, J., & Wilson, K. A. (2017). The Complexities of Counting Fish: Engaging Citizen Scientists in Fish Monitoring. 

Resource managers declared a need for the standardization of river herring monitoring across Maine. The recommended protocol must be adaptable to use in any community, flexible for site specific conditions, comprehensible for volunteer use to ensure data accuracy, and should be updated biannually by regional and state program coordinators and resource managers. 

Lipsky, C., Saunders, R., Stevens, J., O’Malley, M., & Music, P. (2019). Developing sampling strategies to assess the Penobscot River estuary (2010-2013). 

An ensemble of fish capture gear technology can provide a holistic approach to community structure monitoring in estuarine ecosystems.

Nelson, G. A. (2006). A Guide to Statistical Sampling for the Estimation of River Herring Run Size Using Visual Counts. Massachusetts Division of Marine Fisheries Technical Report Series.

The method for estimating run size counts can be determined by using basic statistical concepts, producing reliable estimates, and reviewing statistical designs. This guide recommends a statistical design for community groups to follow.

Wynne, M. P., Clarke, G. A., Saunders, R., Sheehan, T., Collins, M., & Royte, J. (2016). Penobscot I: Monitoring the Penobscot River Restoration Project: Baseline Data to Inform Ecosystem Response.

This slide presentation highlights the objectives that the Penobscot River Restoration Project focused on before and after the removal of dams along the Penobscot river.