fisheries - amazon s3 · 38 changing philosophies of fisheries management as illustrated by the...

Spending Our Fish Dollars

Fisheries

Evolution of Management Philosophies and Fishing Regulations

Vol. 41 • No. 1 • January 2016

Improving Aquatic Invasive Species Detection

Fisheries | www.fisheries.org 1

COLUMNSPRESIDENT'S COMMENTARY

3 Fisheries Down Under Gary Jackson and Ron Essig

POLICY4 Revisiting the Protection/Restoration Debate Thomas E. Bigford

COOL FISH10 No Damsel in Distress Abigail J. Lynch

JOURNAL REVIEWS – 2014 BEST PAPER WINNERS Sarah Harrison5 Postsmolt Growth More Important than Initial Growth at Sea

5 Cluster Sampling Made Easy

STUDENT ANGLES: 2015 Student Writing Contest 6 WINNER Winter: The Forgotten Study Season Michelle Lavery

8 HONORABLE MENTION My Arctic Alter Ego Karen M. Dunmall

AFS NEWS9 Changes at Fisheries Jeff Schaeffer

12 AFS Position Paper and Policy on Mining and Fossil Fuel Extraction

Robert M. Hughes, Felipe Amezcua, David M. Chambers, Wesley M. Daniel, James S. Franks, William Franzin, Donald MacDonald, Eric Merriam, George Neall, Paulo dos Santos Pompeu, Lou Reynolds, and Carol Ann Woody

ESSAYS AND FEATURES16 DefiningForageSpeciestoPreventaManagementDilemma Konstantine J. Rountos

18 Correlation and Causation in Fisheries and Watershed Management

Ray Hilborn

26 Sampling Design for Early Detection of Aquatic Invasive Species in Great Lakes Ports

Joel C. Hoffman, Joshua Schloesser, Anett S. Trebitz, Greg S. Peterson, Michelle Gutsch, Henry Quinlan, and John R. Kelly

38 Changing Philosophies of Fisheries Management as Illustrated by the History of Fishing Regulations in Wyoming

Frank J. Rahel

Vol. 41 • No. 1 • January 2016

Fisheries

38 Trout caught by fly fisherman at West Thumb, Yellowstone Lake, Yellow-stone National Park, Wyoming, 1897 (Nolan 1983).

55 Flapjack octopus in the genus Opisthoteuthis photographed 330 m below the surface in Monterey Bay. Photo credit: 2013 MBARI.

10 Dusky Damselfish Stegastes adustus. Photo credit: Kevin Bryant.

2 Fisheries | Vol. 41 • No. 1 • January 2016

BOOK REVIEW49 Running Silver: Restoring Atlantic Rivers and their Great

Fish Migrations Reviewed by John F. Kocik

51 AFS ANNUAL MEETING 2016

JOURNAL HIGHLIGHTS52 Transactions of the American Fisheries Society Volume 144, Number 6, November 2015

53 CALENDAR

BACK PAGE55 Year in Review: 2015 Natalie Sopinka

Fisheries (ISSN 0363-2415) is published monthly by the American Fisheries Society; 5410 Grosvenor Lane, Suite 110; Bethesda, MD 20814-2199 © copyright 2016. Periodicals postage paid at Bethesda, Maryland, and at an additional mailing office. A copy of Fisheries Guide for Authors is available from the editor or the AFS website, www.fisheries.org. If request-ing from the managing editor, please enclose a stamped, self-addressed envelope with your request. Republication or systematic or multiple repro-duction of material in this publication is permitted only under consent or license from the American Fisheries Society. Postmaster: Send address changes to Fisheries, American Fisheries Society; 5410 Grosvenor Lane, Suite 110; Bethesda, MD 20814-2199.

Fisheries is printed on 10% post-consumer recycled paper with soy-based printing inks.

American Fisheries Society • www.fisheries.org

EDITORIAL / SUBSCRIPTION / CIRCULATION OFFICES5410 Grosvenor Lane, Suite 110•Bethesda, MD 20814-2199(301) 897-8616 • fax (301) 897-8096 • [email protected]

The American Fisheries Society (AFS), founded in 1870, is the oldest and largest professional society representing fisheries scientists. The AFS promotes scientific research and enlight-ened management of aquatic resources for optimum use and enjoyment by the public. It also encourages comprehensive education of fisheries scientists and continuing on-the-job training.

AFS OFFICERSPRESIDENTRon Essig

PRESIDENT-ELECTJoe Margraf

FIRST VICE PRESIDENTSteve L. McMullin

SECOND VICE PRESIDENTJesse Trushenski

PAST PRESIDENTDonna L. Parrish

EXECUTIVE DIRECTORDoug Austen

FISHERIES STAFFSENIOR EDITORDoug Austen

DIRECTOR OF PUBLICATIONSAaron Lerner

MANAGING EDITORSarah Harrison

CONTRIBUTING EDITORSSarah FoxBeth Beard

CONTRIBUTING WRITERNatalie Sopinka

EDITORSCHIEF SCIENCE EDITORSJeff SchaefferOlaf P. Jensen

SCIENCE EDITORSKristen AnsteadMarilyn “Guppy” Blair Jim BowkerMason BryantSteven R. ChippsKen CurrensAndy DanylchukMichael R. DonaldsonAndrew H. FayramStephen FriedLarry M. GigliottiMadeleine Hall-ArborAlf HaukenesJeffrey E. HillDeirdre M. KimballJeff KochJim LongDaniel McGarveyJeremy PrittRoar SandoddenJesse TrushenskiUsha Varanasi Jeffrey WilliamsBOOK REVIEW EDITORFrancis Juanes

ABSTRACT TRANSLATIONPablo del Monte-Luna

ARCHIVE EDITORMohammed Hossain

DUES AND FEES FOR 2016 ARE:$80 for regular members, $20 for student members, and $40 for retired members.

Fees include $19 for Fisheries subscription.

Nonmember and library subscription rates are $191.

Fisheries

Air photo of Imperial Metals' Mount Polley, BC, earthen tailings dam failure on August 4, 2014. The breached dam is in the upper center below the drained tailings pond. The green water and suspended sediments in the lower right show a portion of the pond as it drained into upper Polley Lake before exiting into Hazletine Creek and Quesnel Lake. See Hughes et al. (pages 8-11) for more regarding the hazards of mining and fossil fuel extraction. Photo credit: Cariboo Regional District. British Columbia.

COVER


were on a more ad hoc basis with individuals electing to attend for personal interest reasons.

The ASFB (asfb.org.au) was founded in 1971 and currently has about 325 members including 93 students. As of this writing, there are 34 AFS members from Australia, with most also belonging to ASFB. Administrative work like membership, meeting arrangements, website, and newsletter that were formerly done by volunteers is mostly now done for ASFB under contract. There are two elected representatives from each of the seven Australian states and territories, two from New Zealand, and two more representing the student membership. The ASFB has active Threatened Fishes and Alien Fishes committees, plus several committees that have been active in the past including Stock Assessment, Fish Passageway, and Recreational Fishing. Similar to AFS, all that is needed for such groups to thrive are 1-2 champions to lead them and make them work.

Fisheries Down UnderGary Jackson ([email protected]) and Ron Essig

COLUMNPRESIDENT'S COMMENTARY

AFS President Ron [email protected]

AFS is a lot more than a United States fisheries society. Of course, North America has been at the forefront, but AFS has members from about 60 countries. AFS has been a leader in the World Fisheries Council that organizes the World Fisheries Congress held every four years, with the next being May 23-27, 2016, in Busan, South Korea (wcfs.fisheries.org/7th-world-fisheries-congress-2016-busan-korea). In recent years, AFS has established Memoranda of Understanding (MOU) with the fisheries societies in Brazil, the British Isles, China, Japan, and South Korea for officer exchange programs, where the host society funds lodging and meeting registration for the society president visiting their annual meeting. A similar MOU was just approved with the Australian Society for Fish Biology (ASFB), whereby the ASFB President Gary Jackson attended the 2015 AFS meeting in Portland and AFS President Ron Essig attended the 2015 ASFB conference in Sydney. There have been limited officer exchanges between the two societies in the past, but they Continued on p. 54


Revisiting the Protection/Restoration Debate

from unwise harvest or pressure from non-fishing threats simply makes ecological and economic sense. In the habitat world, the bottom line is that protecting a necessary hydrologic regime or spawning bed is light years easier than attempting to build one.

These options are clear and logical. Why, then, do we all too often practice poor hygiene with fish, to continue the metaphor? Why do we settle for expensive restoration when our wallets argue otherwise? Why don’t we invest more in protecting species from known threats rather than hoping our technical skills will enable us to restore them after their health has been severely compromised? And why is protection deemed to be a bad word to many sectors, including those, such as fishers, who gain from marine protected areas with fishing closures? Unacceptable answers to those policy questions are the root cause of my latent, fish-related anxiety.

Back to those current events noted at the top. BLM’s Arctic focus was the Jack Wade Creek Restoration Project in the Fortymile Wild and Scenic corridor in Alaska. Its efforts aim to reclaim fish habitats and restore stocks decimated by placer mining in the 1800s for alluvial mineral deposits such as gold. Now, equipped with new policies reflecting the latest reclamation techniques, BLM seeks to reclaim values lost for many decades.

Together, the four actions last fall typify our prospects for improved success across all systems. BLM’s challenges in Alaska, and comparable efforts in river systems in the lower 48, are immense, but perhaps not much more than the entire ecosystems at risk in tropical Palau. BLM’s focus on stream design, construction, and monitoring is comparable to work in marine sanctuaries, where successful management hinges on realistic objectives, adequate funding, and thorough monitoring.

The AFS has dabbled in this arena, but not nearly as much as it could and should. AFS wrote to the National Oceanic and Atmospheric Administration in mid-2015 urging a strong scientific approach to designating new sanctuaries. The Society also wrote to the U.S. Environmental Protection Agency and Congress to support their efforts to clarify wetland rules and regulations implementing the Clean Water Act. While those discussions continue, AFS must apply similar logic to fish, fish stocks, and fish habitat. All are easier to protect than to restore.

I usually end my column with a suggestion. Here I implore each of you to be proactive, to balance your options, to be realistic. We can do better.

Opinions are those of the author and not necessarily AFS. Letters to the Editor are invited.

COLUMNPOLICY

AFS Policy DirectorThomas E. [email protected]

This column coalesced around four actions last October. First, as a frequent partner with the U.S. Bureau of Land Management (BLM), the American Fisheries Society (AFS)was invited to a presentation on the agency’s river restoration efforts, land-use planning, and regional mitigation strategies in the Arctic. Second, President Obama announced plans to pursue two new national marine sanctuaries. Third, and on the same day as the sanctuary action, the U.S. Justice Department revealed a $20.8 billion civil settlement addressing a portion of the environmental harm associated with BP’s Gulf of Mexico oil spill in 2010. Finally, to punctuate the flurry of poignant messages, Palau designated a 193,000 square-mile marine reserve, the sixth largest protected area in the world. From the Arctic to Palau, there are policies to debate and lessons to learn. You’ll sense some anguish in this column, prompted because we absolutely must find better balance in these two extremes—protect and restore.

At the heart of these actions is a choice. The options apply to fish habitat (the traditional bounds) but equally to fish stocks. In both arenas, the options can be clarified by parallels in our personal life. Consider our overall health, where the protective course of action is regular check-ups with timely immunizations and perhaps some vitamin supplements; the restoration approach to our well-being would be reactive, kicking into full gear after our health declines, then relying on prescription medicine and surgery to address serious disease. In dentistry, protection is akin to fluoride, flossing, and brushing, supplemented with regular visits. Restoration is decisive, but not until damage is detected, and we’re fighting cavities, drilling root canals, and perhaps checking our insurance coverage for dentures. Back in the fish world, my two extremes are not nearly so distinct. Regardless of where we draw the lines, it still makes obvious sense to invest in protection rather than settle for restoration.

As noted, this debate often focuses on habitat, but projecting those examples to fish is hardly a stretch. The ecological and economic costs of both options are at the root of the four actions mentioned in my opening paragraph. Protection is a strong decision to defend human values while reducing risks. It’s about minimizing and avoiding. In its simplest form, protecting stocks

Protecting stocks from unwise harvest or pressure from non-fishing threats simply makes ecological and economic sense.

Thomas E. Bigford | AFS Policy Director


WINNER: Best Paper, Marine and Coastal Fisheries

Postsmolt Growth More Important than Initial Growth at Sea

Over the last 20-30 years, the abundance of steelhead Oncorhynchus mykiss in the Keogh River, British Columbia, has declined substantially. Increased marine mortality has been cited as the main reason for this decline, with size at ocean entry appearing to be the main factor determining recruitment. However, in a new study in Marine and Coastal Fisheries, Kevin D. Friedland of the National Marine Fisheries Service and his colleagues found other factors to be affecting marine survival. Through restrospective growth analy-sis using scales from returning steelhead, they found that the growth shortly after smolt transition was a strong predictor of survival to adulthood in the Keogh River from 1977 to 1999. Further stating that this growth may alleviate any initial size differences in smolts, lessening the effect of size of ocean entry. They also found that this growth was closely related to sea surface temperatures during the June-December period following ocean entry. This article is important because it reconfirms that the survival pattern of the Keogh River steelhead population has changed, with sustained growth conditions during summer and fall of the postsmolt year being more important than initial growth at sea.

REFERENCEFriedland, K. D., B. R. Ward, D. W. Welch, and S. A. Hayes. 2014. Postsmolt growth and thermal regime define the marine survival of steelhead

from the Keogh River, British Columbia. Marine and Coastal Fisheries 6:1-11. dx.doi.org/10.1080/19425120.2013.860065

WINNER: Best Paper, Transactions of the American Fisheries Society

Cluster Sampling Made EasyCluster sampling is one of the most common survey designs used in fisheries research to sample fish populations. However, many

researchers don’t take clustering into account when they are analyzing their data, largely due to lack of awareness. When the effect of clustering is ignored, population estimates, results of hypothesis testing, and conclusions drawn from statistical analyses may be incor-rect. In Transactions of the American Fisheries Society, Gary A. Nelson of the Massachusetts Division of Marine Fisheries provides a nice article on the nature of cluster sampling design and common mistakes researchers make when using it. He provides an excellent introduction on how to estimate population attributes and analyze fisheries data collected via cluster sampling.

We loved this paper because it has an important practical application, as well as a thoughtful discussion that provides insight into understanding fish sampling design, making it a must read for any fisheries class.

REFERENCENelson, G. A. 2014. Cluster sampling: a pervasive, yet little recognized survey design in fisheries research. Transactions of the American

Fisheries Society 143(4):926-938. dx.doi.org/10.1080/00028487.2014.901252

Find the 2014 Best Paper winners for the journals, Journal of Aquatic Health, North Ameri-can Journal of Aquaculture, and North American Journal of Fisheries Management, in the December 2015 issue.

JOURNAL REVIEWS – 2014 WINNERS

Sarah Harrison | AFS Managing Editor. E-mail: [email protected]


WINNER

Winter: The Forgotten Study Season

STUDENT ANGLES 2015 Student Writing Contest

Michelle Lavery | MSc Candidate, Cunjak Lab, Canadian Rivers Institute, University of New Brunswick, 10 Bailey Dr., Fredericton, NB, E3B5A3, Canada. E-mail: [email protected]

For many hydrologic regimes of the world, streams and rivers are ice covered for the majority of the year, yet minimal research is conducted during this period compared with the more “researcher-friendly” open-water period. Without a doubt, scientific progress is hampered by the logistical difficulties and high cost associated with conducting “winter” research. (Prowse, 2001 [part II]) It seems as though every winter ecology paper contains some

variant of this sentiment—we know that winter is important, but we’re not crazy enough to study it. As researchers, we’ve built sampling regimes that ignore an entire season because winter is considered harsh and unforgiving. It’s cold, sharp, and sometimes deadly to us, and so we operate under the assumption that the same goes for the creatures we study.

Alas, it is not so. There’s a lot going on under the snow, and even more going on under the ice. For example, Atlantic salmon eggs incubate in the gravel under river ice in Eastern Canada for six frigid, snowy months at water temperatures barely above freezing. They emerge from the gravel during the spring melt period, when ice jams bulldoze forests and water levels climb metres in minutes. These tiny fish are at the mercy of a dynamic and unpredictable season, yet we barely know anything about it.

As a pampered girl from “tropical” Toronto, I never imagined myself riding a snowmobile and hacking through river ice in the middle of the woods. However, through a serendipitous connection, I recently found myself doing both—while pursuing a master’s degree supervised by Richard Cunjak at the Canadian Rivers Institute.

In Eastern Canada’s Miramichi River system, salmon eggs incubate in the gravel riverbed from late October to early May, during which time they experience highly variable winter conditions. In November, air temperatures can drop dramatically overnight (usually to about -20°C or -4°F), causing water to reach its freezing point quickly and inconsistently. As water crystallizes, it can stick to itself and the river substrate, forming anchor ice—a squishy carpet of ice crystals on the riverbed. If this ice forms on top of salmon nests (or “redds”), it can block water flow through the gravel and alter the temperature and dissolved oxygen levels surrounding the developing eggs.

Once full ice cover forms and surface runoff is locked up in the snowpack, long-residence groundwater may be the major contributor to river discharge. “Long-residence” groundwater refers to water that has spent a considerable amount of time in an aquifer deep underground. Consequently, it is often warmer than the surface water in the winter, and can have significantly lower dissolved oxygen concentrations (since it has not been recently aerated). As this groundwater seeps through the river substrate, the conditions in salmon redds can change dramatically. Depending on the size of the seep, eggs may develop faster due to warmer water temperatures and require more oxygen

to sustain this accelerated rate of development. However, the oxygen-poor groundwater is usually unable to meet their biological demands. Without enough oxygen, these eggs may die or experience “sub-lethal” effects—consequences that may impact their survival later in life as free-swimming fish. These may include stunted growth or developmental deformities that may impair gas exchange, swimming ability, neurological function, etc.

During the spring melt period, silt and clay can be eroded into rivers by the surface runoff water. Depending on the grain size, these sediments may clog the egg membrane and prevent diffusion of oxygen to the embryo, effectively suffocating the fish. Furthermore, as ice breaks up and moves out of rivers, scour along the riverbed may significantly disturb the gravel and damage the embryos underneath.

It is hard to believe, after considering all of the variation inherent in winter and its potential effects on one life stage of one species in one type of habitat, that winter goes largely unnoticed in the scientific literature. It is, certainly, a challenging season to research. I’ve had my fair share of winter mishaps, including digging a snowmobile out of a slush puddle for three


hours, miscalculating ice thickness (not ideal!), hypothermic near-misses, and tethering myself to a tree during the spring melt. However, if we can get past our numb fingers and dripping noses, there’s a whole season waiting to be studied. One could argue that winter research is the last true frontier of freshwater ecology—there are so many unknowns to explore, and so many questions left unanswered. It might not be a “researcher-friendly” season, but it’s definitely exciting! Plus, who doesn’t love a good mid-river snowball fight?

REFERENCEProwse, T. D. 2001. River-ice ecology. 2: Biological aspects. Journal

of Cold Regions Engineering 15:17-33.

RELATED MATERIALSCunjak, R. A., T. D. Prowse, and D. L. Parrish. 1998. Atlantic Salmon

(Salmo salar) in winter: “the season of parr discontent”? Cana-dian Journal of Fisheries and Aquatic Sciences 55(1):161-180.

Flanagan, J. J. 2003. The impacts of fine sediments and variable flow regime on the habitat and survival of Atlantic Salmon (Sal-mo salar) eggs. Master’s thesis: University of New Brunswick, Fredericton.

Greig, S. M., D. A. Sear, and P. A. Carling. 2005. The impact of fine sediment accumulation on the survival of incubating salmon progeny: Implications for sediment management. Science of the Total Environment 344(1-3):241-258.

Kane, T. R. 1988. Relationship of temperature and time of initial feed-ing of Atlantic Salmon. Progressive Fish-Culturist 50:93-97.

Louhi, P., M. Ovaska, A. Maki-Petays, J. Erkinaro, T. Muotka, and J. Rosenfeld. 2011. Does fine sediment constrain salmonid alevin development and survival? Canadian Journal of Fisheries and Aquatic Sciences 68(10):1819-1826.

Malcolm, I. A., S. M. Greig, A. F. Youngson, and C. Soulsby. 2008. Hy-porheic influences on salmon embryo survival and performance. Pages 1-24 in D. A. Sear and P. DeVries, editors. Salmonid spawn-ing habitat in rivers: physical controls, biological responses, and approaches to remediation. American Fisheries Society, Sympo-sium 65, Bethesda, Maryland.

Malcolm, I. A., C. Soulsby, A. F. Youngson, D. M. Hannah, I. S. McLaren, and A. Thorne. 2004. Hydrological influences on hyporheic wa-ter quality: implications for salmon egg survival. Hydrological Processes 18(9):1543-1560.

Malcolm, I. A., C. Soulsby, A. F. Youngson, and D. M. Hannah. 2005.Catchment-scale controls on groundwater-surface water inter-actions in the hyporheic zone: implications for salmon embryo survival. River Research and Applications 21(9):977-989.

Malcolm, I. A., C. Soulsby, A. F. Youngson, and D. Tetzlaff. 2008b. Fine scale variability of hyporheic hydrochemistry in salmon spawning gravels with contrasting groundwater-surface water interactions. Hydrogeology Journal 17(1):161-174.

Malcolm, I. A., A. F. Youngson, and C. Soulsby. 2003. Survival of sal-monid eggs in a degraded gravel-bed stream: effects of ground-water-surface water interactions. River Research and Applica-tions 19(4):303-316.

Prowse, T. D. 2001. River-ice ecology. 1: hydrologic, geomorphic, and water-quality aspects. Journal of Cold Regions Engineering 15:1-16.

Soulsby, C., A. F. Youngson, H. J. Moir, and I. A. Malcolm. 2001. Fine ediment influence on salmonid spawning habitat in a lowland agricultural stream: a preliminary assessment. Science of the To-tal Environment 265(1-3):295-307.

mitpress.mit.edu

An analysis of how responsive governance has shaped the evolution of global fi sheries in cyclical patterns of depletion and rebuilding dubbed the “manage-ment treadmill.”

The MIT Press


HONORABLE MENTION

My Arctic Alter EgoKaren M. Dunmall | [email protected]

As the sun slowly rises, we race across the frozen Mackenzie River Delta in a helicopter. The landscape is black and white; the twilight hoards the colours to paint the sky. We have donned our many layers

of wool and feathers to ward off the extreme cold. Our food is in a cooler with a hot water bottle to keep it from freezing. As we fly over the icy maze of lakes and channels, I am mentally shifting. I am a mom that is headed into an unforgiving, frozen world. Rather than coordinating the usual hockey practices, birthday parties, and playdates, today my responsibility is to get my research team back home safely. If we get data too, that is a bonus. The helicopter is my phone booth.

I am the only person in the world that is researching salmon colonizations in the Canadian Arctic. While salmon are getting caught in higher abundances in subsistence harvests throughout the Canadian Arctic, it is not known where they may successfully colonize and if they will compete with local fish species. Even in the darkest days of the Arctic winter, there are rare places that do not freeze. In the summer, these places blend in with the landscape; in winter, their camouflage is lost and the dark, flowing water is starkly contrasted against the sparkling snow-covered trees and frosty willows. These are spawning locations for Dolly Varden Salvelinus malma, a cold-tolerant native char with similar habitat requirements to salmon. The goal is to determine if those places may also be viable for spawning vagrant salmon. Last winter, we deployed devices to record water temperature in these locations. Today, we are headed back to retrieve the data and redeploy the devices.

The helicopter lands and we emerge into waist-deep snow. We clumsily move toward the riverbank. The temperature loggers are in metal pipes that have been driven into the river bottom. I turn on a metal detector and sweep the wand back and forth as I walk along a section of shallow, open water. I wait for the hum to turn into a squeal, pinpointing the location

of the pipe. Another team member kneels and twists off the lid, holding up the loggers for all to see. I pull a new lid, with loggers attached, out of my chest waders, and screw it onto the pipe. The extra room in the waders from the male-inspired design provides me a useful storage pouch. As we wade back toward the helicopter, our wet boots and waders accumulate layers of ice. I am teetering on an extra four inches by the time we reach the skids. As the helicopter blades start to spin, I suck on a frozen M&M candy and load up the coordinates for the next stop. We repeat this process until the sun sinks low on the horizon once again.

As we race back to the hanger to land before dark, I rub the window in a circle with my mitten to clear the frost and pause to own this moment of success. It hurts to leave my family and winter Arctic fieldwork carries significant risks. However, I look forward to the squeals that signal mommy’s return and I know we will soon settle back into the everyday routine. I am teaching my kids by example that it is ok to follow their dreams. Eventually they will understand that a challenging journey reaps the highest rewards.

To follow the Arctic Salmon research, “like” it on Facebook: facebook.com/arcticsalmon.

Karen Dunmall is a Ph.D. student at the University of Manitoba, working in collaboration with Fisheries and Oceans Canada. She is expertly guided by supervisors at both institutions. She gratefully acknowledges receiving a W. Garfield Weston Foundation Award for Northern Research, an NSERC Alexander Graham Bell Canada Graduate Scholarship, the American Fisheries Society 2014 J. Frances Allen Scholarship, and the University of Manitoba Roger Evans Memorial Scholarship. Her research is funded by Fisheries and Oceans Canada, University of Manitoba, Government of the Northwest Territories through the Northwest Territories Cumulative Impact Monitoring Program (Project # 00142), Fisheries Joint Management Committee, Gwich’in Renewable Resources Board, Gwich’in Land Use Planning Board, and the Sahtu Renewable Resources Board.

STUDENT ANGLES 2015 Student Writing Contest

Sunrise over the Mackenzie River Delta, NWT. Photo credit: K. Dunmall.

Visible open water areas in Fish Creek, NWT. Photo credit: K. Dunmal.


Changes at Fisheries Jeff Schaeffer

It is with mixed emotions that this month we bid farewell to Sarah Gilbert Fox, our long-standing man-aging editor at Fisheries. Fox is leaving us to return to the world of creative writing (rumor has it that her second Great American Novel is forthcoming), and a broad range of personal endeavors that were on hold for far too long. We are sad to see her go, but filled with joy that she is going back to her one great passion.

We owe Fox a huge debt, because for quite some time she was the face of Fisheries, and facilitated its continuing evolution into what is clearly a modern, well-designed professional society magazine. She did this almost entirely through force of personality, by knowing everyone in AFS, and by forging deep, personal connections with people across the Society. Her passion was pervasive, her touch deft, and her commitment to AFS was deep. We will miss her, but wish her the best in her new life. And of course, many of us will remain connected to her via social media and likely Amazon pre-orders of her new books.

Our new managing editor will be Sarah Harrison. She is an experienced science writer and has taken on an ever-expanding role. The most recent issue of Fisheries is largely her work. Harrison brings some new and welcome skills to our group that we hope to use in new and creative ways on the magazine side of publication. Please welcome her as she grows her new role.

Perhaps the best thing that Sarah Gilbert Fox brought to AFS was her sense of cognition in the face of mayhem. Fisheries is a monthly publication, and the week prior to printing is always a source of stress. The features and articles need to be perfect and accurate, authors are submitting changes to page proofs, the printed version has to be eye-catching, and it all has to fit into a precisely defined print space. It is a lot like moving to a new place you have never been before, and wondering how all the furniture and oddly-shaped boxes can possibly fit into the back of the truck. Inevitably, there would be a host of last minute changes and corrections. Fox would often stay up all night fixing our mistakes, but every issue came together on time despite a myriad of last-minute, eleventh-hour tweaks. And the content kept getting better and better, especially when we simply unleashed her vast creative energy.

Although Fox is leaving, there is one connection with AFS that will remain inviolate. Whenever one of us would mess things up so badly, such that it required her to stay up all night, we would get an email or phone message that stated, “You owe me one dinner.” This was, of course, a kindhearted and humorous way of letting you know that you had created a real mess that took great effort to clean up. Sarah Harrison never owed any dinners, Olaf Jensen (co-chief science editor) owes two, and my total was somewhere between 118 and 137 (19 dinners are still in mediation). Fox, you will be my friend forever, and you will get every single one of those dinners.

HONORABLE MENTION

My Arctic Alter EgoKaren M. Dunmall | [email protected]

THE VALUE OF FISH CULTUREThe value of fish culture and the extent and value of the fishery industries are well known to the members of this body and

to those directly interested, but with the great mass of our population they are' not properly appreciated.

Prof. Baird has truly said (Report, 1890): "It may be safely stated that, as a source of animal food to man, the sea is the great fountain-head, and that without this resource the supply of such food would be comparatively limited and far inferi-or to the demand of the various populations of the globe. In the much greater population of ocean to land this reservoir of food is practically inexhaustible; and not only do the people living near its shores find a daily supply for consumption in a fresh state, but, by proper methods of preparation and preservation, the product of the sea can be fitted for long-continued keeping and for transportation to distant markets, where fishing is difficult, or into the interior, where it is impracticable."

John Gay & Wm P. Seal (1890) The past and Present of Fish Culture, with an Inquiry as to what may be Done to Further Promote and Develop the Science, Transactions of the American Fisheries Society, 19:1, 73

FROM THE ARCHIVES

AFS NEWS


COLUMNCOOL FISH

No Damsel in Distress I’m not sure how many people can trace

their career origin back to a specific instance, but I can.

I was sitting in a concrete block of a building, an old military installation that had been retrofitted to be an auditorium-style classroom. Students were scattered across the carpeted steps of the room. It was hard to tell if the carpet was damp or just dingy from years of exposure to the salty sea air. There was a big chalkboard in front of us; the professor started a list of marine organisms. We would draw numbers, we were told, to pick an organism from the list on the board. Our choice would be the focus of our independent research projects.

I was an undergraduate student in the long-running University of Virginia Marine Biology Study Abroad program. We were on San Salvador, a tiny, 65-square mile island at the easternmost edge of the Bahamas. San Sal, purportedly, was the first glimpse Christopher Columbus had of the new world.

Sitting in that auditorium, we’d been through about half of our program, with lectures in the morning and evening and snorkeling most of the day. We soaked up everything we could, hoping our brains could hold as much information as a sea sponge could hold water. Now it was time for us to take the science training-wheels off and apply all that we had learned to our very own research projects.

Excited and nervous, I drew my number—a woefully high number—oh no!! I had made the mistake of setting my hopes on studying the

Abigail J. Lynch | National Climate Change and Wildlife Science Center, U.S. Geological Survey, 12201 Sunrise Valley Drive, MS-516, Reston, VA 20192. E-mail: [email protected]


Dusky Damselfish Stegastes adustus—by far, the coolest thing on the list—it would surely be selected by the time my number was called.

Amid a rainbow of colorful reef fish, this drab little fish stands out. On one of our first snorkeling trips, I noticed these fish on the reef bed nibbling on algae from a well-defined patch of reef, which they appeared to be guarding. Turns out, they were. Dusky Damsels are essentially algal gardeners. In tending their territories, Dusky Damsels have a measurable effect on the benthic community. They foster highly productive gardens with higher biomass and greater algal diversity than at locations outside their territories. By defending their gardens from other herbivores, they ensure a limitless food source for themselves (their consumption never exceeds primary production).

As a curious snorkeler, I found out fast that the territoriality of Dusky Damsels is pretty astounding—when I dove down to take a look at one of these garden plots, I got an assertive tap on the mask from the garden’s owner! I was shocked and charmed all at once. This was no damsel in distress. This plucky little fish had me hooked.

So, as I sat still waiting for my number to be called, Dusky Damselfish remained written on the board in big block letters, unclaimed. Ghost crab—gone. Eel grass—gone. Even Halimeda algae—gone. None of the prior students picked my fish. I was floored. Who wouldn’t want to study damselfish? They are so cool!

Until that moment, it had never occurred to me that others didn’t share my passion for all that is piscine. I had just assumed that everyone thought that fish were as fascinating as I did. In the end, I was able to select Dusky Damsels and my small, highly un-scientific examination of their territoriality response to predator and competitor replicas became my turning point.

Looking back on it now, I’m not sure why all of this came as such a surprise to me. I suppose, at the time, I hadn’t considered that my interests were differentiating. I hadn’t realized that I was unintentionally developing a professional path. I just loved every minute of that project. When I discovered that I could make a career of this, I wouldn’t have had it any other way.

ACKNOWLEDGMENTS

I thank the instructors and the student cohort of the 2003 Marine Biology and Coral Reef Ecology in San Salvador program, in particular Professor Fred Diehl (Doc). The excitement and enthusiasm of this program inspired my career path and still inspires me today.

Abby Lynch hand-made a damselfish cake topper for her wedding cake. She and her husband recently celebrated their first anniversary. Photo credit: Bret Thacker.

Dusky Damselfish Stegastes adustus. Photo credit: Kevin Bryant.

Juvenile Dusky Damselfish Stegastes adustus. Photo credit: Kevin Bryant.


Robert M. HughesAmnis Opes Institute and Department of Fisheries & Wildlife, Oregon State University, Corvallis, OR 97333. E-mail: [email protected]

Felipe AmezcuaInstituto de Ciencias del Mar y Limnologia, Universidad Nacional Autónoma de Mexico, Mazatlán, Sinaloa, México

David M. ChambersCenter for Science in Public Participation, Bozeman, MT

Wesley M. DanielDepartment of Fisheries and Wildlife, Michigan State University, East Lansing, MI

James S. FranksGulf Coast Research Laboratory, University of Southern Mississippi, Ocean Springs, MS

William FranzinLaughing Water Arts & Science Inc., Winnipeg, Manitoba, Canada

Donald MacDonaldMacDonald Environmental Sciences Limited, Nanaimo, British Columbia, Canada

Eric MerriamDivision of Forestry, Natural Resources and Design, West Virginia University, Morgantown, WV

George NeallRetired Mining Engineer, Fulks Run, VA

Paulo dos Santos Pompeu Departmento de Biologia, Universidade Federal de Lavras, Lavras, Minas Gerais, Brazil

Lou ReynoldsFreshwater Biology Team, U.S. Environmental Protection Agency Region III, Wheeling, WV

Carol Ann WoodyCenter for Science in Public Participation, Anchorage, AK

Following a four-year period of writing, member comment, and multiple revisions, the AFS Position Paper and Policy on Mining and Fossil Fuel Extraction was approved unanimously by the membership at the Society’s annual business meeting August 19, 2015, in Portland, Oregon. The entire document can be read at fisheries.org/policy_statements; a brief summary follows.

AFS Position Paper and Policy on Mining and Fossil Fuel Extraction

AFS NEWS

TECHNICAL BACKGROUND

Mining (hard-rock, aggregate, deep, and surface) and fossil fuel (coal, oil, gas) extraction have the potential to significantly alter aquatic ecosystem structure and function. Adverse impacts on water quality, hydrology, physical habitat structure, aquatic biota, and fisheries include elimination and contamination of receiving waters (USEPA 2011; USEPA 2014); significantly altered algal, macroinvertebrate, and fish assemblages (e.g., Pond et al. 2008; Lavoie et al. 2012; Daniel et al. 2014; Figure 1); impairments of aquatic-dependent wildlife (USEPA 2011; USEPA 2014); and climate change (Hansen et al. 2013).

For example, even at low concentrations, mining-associated contaminants, such as copper, impair salmonid olfactory function (McIntyre et al. 2008), thereby increasing predation susceptibility (McIntyre et al. 2012); alter salmonid migratory behavior (Sprague et al. 1965; Lorz and McPherson 1976); increase disease susceptibility (Baker et al. 1983); and reduce growth (Marr et al. 1996).

Despite predicted compliance of permit conditions, many operating metal mines have violated water quality criteria (Kuipers et al. 2006). Those permit conditions, or applicable regulations, are minimum requirements and typically do not represent best management practices. Also, the applicable regulations rarely account for the cumulative effects of pollution from multiple mines.

Under the General Mining Act of 1872 in the United Sates, federal law transfers metal wealth from the public to mining companies, and shifts clean-up liability from those companies to taxpayers. The half-million abandoned hard-rock mines in the United States could cost US$72-240 billion to rehabilitate (USEPA 2000; NRC 2005), and the over 100 abandoned metal mines in Quebec are estimated to cost over US$600 million to remediate (Hamilton et al. 2015). In both cases, the majority of those costs will fall on taxpayers (Woody et al. 2010; Chambers et al. 2012; Hamilton et al. 2015). In addition, those estimates do not include rehabilitation of the newer, larger mines being


developed in more inhospitable environments, nor the costs of spills, failures, and accidents. Such a spill into the Animas River occurred on August 5, 2015, when U.S. Environmental Protection Agency contractors were assessing leaks of toxic metals in the vicinity of the abandoned Red, Bonita, and Gold King mines near Silverton, Colorado, and broke the debris dam of a holding pond (Vox 2015). That spill sent 12 million liters of contaminated water downstream and into the states of New Mexico and Utah via the San Juan River; the costs of fish kills, water monitoring, clean-up, and loss of water use by recreationalists, irrigators, and other users have been estimated in the tens of millions of dollars. The Mt. Polley tailings pond failure on August 4, 2014, which spilled 14 million m3 of metals-contaminated water and sediments into Hazeltine Creek and Polley and Quesnel Lakes in central British Columbia, has been estimated to cost at least $500 million (US) to rehabilitate (Uechi 2014). Because of various economic factors, the numbers of serious and very serious tailings dam failures such as these have increased since 1960 (Bowker and Chambers 2015).

Surface mining temporarily eliminates surface vegetation and permanently changes topography and hydrology, as with mountain-top-removal-valley-fill (MTRVF) coal mines (Fritz et al. 2010; USEPA 2011). Reclaimed surface mines create a leach bed for ions producing toxic conductivity concentrations (Pond et al. 2008), whereas altered hydrology produces flashy flows similar to those in urban areas (Ferrari et al. 2009; USEPA 2011). Underground mines produce acid mine drainage that can eliminate most aquatic life across extensive regions or alkaline mine drainage that alters ionic balance of freshwater ecosystems (USEPA 1995; USEPA 2000). Instream and gravel bar aggregate mining and dredging can alter channel morphology and increase bed and bank erosion, which can reduce riparian vegetation and impair downstream aquatic habitats (Kondolf 1994).

Oil and gas wells and product transport can cause devastating spills in freshwater and marine ecosystems (e.g., AP 2012; Amezcua-Linares 2013; Keller 2015). Hydraulic fracturing to extract residual oil and gas can contaminate

groundwater and alter surface water ecosystems (Entrekin et al. 2011; Weltman-Fahs and Taylor 2013). Fish biodiversity and macroinvertebrate taxa richness were negatively correlated with the number of well pads per catchment and total mercury concentrations in crayfish, and Brook Trout Salvelinus fontinalis were positively correlated with well pads per catchment (Grant et al. 2015). Smith et al. (2012) concluded that fewer than one well pad per 3 km2 and 3 ha per pad were needed to minimize damage to Brook Trout populations. Bamberger and Oswald (2012) and Webb et al. (2014) documented a wide range of health effects as a result of exposure to fracking fluids and gases. McKenzie et al. (2012) reported cumulative cancer risks of 10 per million for residents living <800 m from a gas well. Stacey et al. (2015) and Cil (2015) associated lowered infant health with proximity of drinking water wells to gas wells. The casings and grouting of abandoned oil and gas wells should be expected to eventually leak and contaminate surface and ground water (Dusseault et al. 2000). In addition, fossil fuel combustion is fundamentally altering the global climate, sea levels, and ocean chemistry (e.g., Orr et al. 2005; Dai 2013; Hansen et al. 2013).

Catastrophic mine tailings failures have killed hundreds of thousands of fish and hundreds of people, and contaminated tens to thousands of river kilometers (USEPA 1995; Chambers and Higman 2011; WISE 2011). Oil and gas wells are exempted from regulation by several U.S. laws, despite considerable evidence of their detrimental effects on surface and ground water (Allen et al. 2011; Amezcua-Lineares 2013; Keller 2015).

PROPOSED AFS POLICY

Mines and wells should only be developed where, after weighing multiple costs, benefits, beneficiaries, and liabilities, they are considered the most appropriate use of land and water by affected publics, can be developed in an environmentally responsible manner, benefit workers and affected communities, and are appropriately regulated. Because of substantial widespread adverse effects of mining and wells on aquatic ecosystems and related human communities, fossil fuel

Figure 1. Percent generally intolerant fish individuals as a function of mine density for the conterminous USA (n = 33,538). Mines include coal, hard rock, uranium, and aggregate mines.


combustion effects on global climate, and enormous unfunded reclamation costs for abandoned extraction sites, the American Fisheries Society (AFS) recommends substantive changes in how North American governments conduct environmental assessments and permit, monitor, and regulate mine and fossil fuel development. In particular, AFS recommends that:

1. Following a formal environmental impact assessment, the affected public should be involved in deciding whether a mine or well is the most appropriate use of land and water, particularly relative to the need to preserve ecologically and culturally significant areas.

2. Mine or well development should be environmentally responsible with regulation, treatment, monitoring, and sureties sufficient for protecting the environment in perpetuity.

3. Baseline ecological and environmental research and monitoring should be conducted in areas slated for mining and fossil fuel extraction before, during, and after development so that the effects of those industries can be assessed in an ecologically and statistically rigorous manner, and the resulting data should be made publicly available.

4. This policy and related research should help inform the process of responsible resource development for mining and fossil fuel extraction, and should guide the implementation of the precautionary principle for those sectors.

5. A formal risk assessment of the cumulative atmospheric, aquatic, and oceanic effects of continued fossil fuel extraction and combustion should be conducted and reported to the public.

6. A formal risk assessment of the cumulative aquatic and oceanic effects of continued hard rock and aggregate extraction and metals smelting should be conducted and reported to the public.

ACKNOWLEDGMENTS

This manuscript was improved by review comments from Leanne Roulson. The views expressed in this article are those of the authors and do not necessarily reflect the views or policies of the U.S. Environmental Protection Agency. We thank the National Fish Habitat Partnership and funding from the U.S. Fish and Wildlife Service and the U.S. Geological Survey for supporting development of the data used in Figure 1.

REFERENCESAllen, L., M. J. Cohen, D. Abelson, and B. Miller. 2011. Fossil fuels and

water quality. Pages 73-96 in P. H. Gleick, L. Allen, J. Christian-Smith, M. J. Cohen, H. Cooley, M. Heberger, J. Morrison, M. Pala-niappan, and P. Schulte, editors. The world’s water, volume 7. Island Press, Washington, DC.

Amezcua-Linares, F., F. Amezcua, and B. Gil. 2013. Effects of the Ix-toc I oil spill on fish assemblages in the southern Gulf of Mexico. Pages 209-235 in: B. Alford, M. Peterson, and C. Green, editors. Impacts of oil spill disasters on marine fisheries in North Ameri-ca. Taylor & Francis, New York.

AP (Associated Press). 2012. Exxon increases estimate of Yellowstone River oil spill by 50%. Website. Available: billingsgazette.com/news/local/exxon-increases-estimate-of-yellowstone-river-oil-spill-by/article_e3f0de2e-f931-50e8-9678-c5f230c9e00d.html. (August 2015).

Baker, R. J., M. D. Knittel, and J. L. Fryer. 1983. Susceptibility of Chi-nook Salmon, Oncorhynchus tshawytscha (Walbaum), and Rain-bow Trout, Salmo gairdneri Richardson, to infection with Vibrio

anguillarum following sublethal copper exposure. Journal of Fish Diseases 6:267–275.

Bamberger, M., and R. E. Oswald. 2012. Impacts of gas drilling on human and animal health. New Solutions 22:51-77.

Bowker, L. N., and D. M. Chambers. 2015. The risk, public liability, and economics of tailings storage facility failures. Available: www.earthworksaction.org/files/pubs-others/BowkerChambers-RiskPublicLiability_EconomicsOfTailingsStorageFacility%20Failures-23Jul15.pdf. (August 2015).

Chambers, D. M., and B. Higman. 2011. Long term risks of tailings dam failure. Center for Science in Public Participation, Bozeman, Montana.

Chambers, D., R. Moran, L. Trasky, S. Bryce, L. Danielson, L. Fulker-son, J. Goin, R. M. Hughes, J. Konigsberg, R. Spies, G. Thomas, M. Trenholm, and T. Wigington. 2012. Bristol Bay’s wild salmon ecosystems and the Pebble Mine: key considerations for a large-scale mine proposal. Wild Salmon Center and Trout Unlimited, Portland, Oregon.

Cil, G. 2015. Effects of behavioral and environmental factors on in-fant health. Doctoral dissertation. Department of Economics, University of Oregon, Eugene, Oregon.

Dai, A. 2013. Increasing drought under global warming in observa-tions and models. Nature Climate Change 3:52-58.

Daniel, W. M., D. M. Infante, R. M. Hughes, P. C. Esselman, Y.-P. Tsang, D. Wieferich, K. Herreman, A. R. Cooper, L. Wang, and W. W. Taylor. 2014. Characterizing coal and mineral mines as a regional source of stress to stream fish assemblages. Ecological Indica-tors 50:50-61.

Dusseault, M. B., M. N. Gray, and P. A. Nawrocki. 2000. Why oil wells leak: cement behavior and long-term consequences. Proceed-ings of the International Oil and Gas Conference and Exhibition in China. Society of Petroleum Engineers DOI:10.2118/64733-MS.

Entrekin, S., M. Evans-White, B. Johnson, and E. Hagenbuch. 2011. Rapid expansion of natural gas development poses a threat to surface waters. Frontiers in Ecology and the Environment 9:503-511.

Ferrari, J. R., T. R. Lookingbill, B. McCormick, P. A. Townsend, and K. Eshleman. 2009. Surface mining and reclamation effects on flood response of watersheds in the central Appalachian Plateau region. Water Resources Research 45(4):1–11.

Fritz, K. M., S. Fulton, B. R. Johnson, C. D. Barton, J. D. Jack, D. A. Word, and R. A. Burke. 2010. Structural and functional character-istics of natural and constructed channels draining a reclaimed mountaintop removal and valley fill coal mine. Journal of the North American Benthological Society 29:673-689.

Grant, C. J., A. B. Weimer, N. K. Marks, E. S. Perow, J. M. Oster, K. M. Brubaker, R .V. Trexler, C. M. Solomon, and R. Lamendella. 2015. Marcellus and mercury: assessing potential impacts of un-conventional natural gas extraction on aquatic ecosystems in morthwestern Pennsylvania. Journal of Environmental Science and Health, Part A 50:482-500.

Hamilton, P. B., I. Lavoie, S. Alpay, and K. Ponader. 2015. Using dia-tom assemblages and sulfur in sediments to uncover the effects of historical mining on Lake Arnoux (Quebec, Canada): a ret-rospective of economic benefits vs. environmental debt. Fron-tiers in Ecology and Evolution [online serial] 3:99. DOI: 10.3389/fevo.2015.00099.

Hansen, J., P. Kharecha, and M. Sato. 2013. Climate forcing growth rates: doubling down on our Faustian bargain. Environmental Research Letters 8:1-9.

Keller, J. 2015. The maddening silver lining to BP’s $18.7 billion pen-alty. Pacific Standard. Available: www.psmag.com/politics-and-law/how-come-bp-gets-to-treat-fines-like-business-expenses-but-i-cant-even-get-out-of-this-parking-ticket. (August 2015).

Kondolf, G. M. 1994. Geomorphic and environmental effects of in-stream gravel mining. Landscape and Urban Planning 28:225–243.

Kuipers, J. R., A. S. Maest, K. A. MacHardy, and G. Lawson. 2006. Comparison of predicted and actual water quality at hardrock mines: the reliability of predictions in environmental impact statements. Kuipers and Associates, Butte, Montana.

Lavoie, I., M. Lavoie, and C. Fortin. 2012. A mine of information: benthic algal communities as biomonitors of metal contamina-tion from abandoned tailings. Science of the Total Environment 425:231-241.

Lorz, H. W., and B. P. McPherson. 1976. Effects of copper or zinc in fresh water on the adaptation to seawater and ATP-ase activ-ity and the effects of copper on migratory disposition of Coho Salmon. Journal of the Fisheries Research Board of Canada 33:2023-2030.


Marr, J. C. A., J. Lipton, D. Cacela, J. A. Hansen, H. L. Bergman, J. S. Meyer, and C. Hogstrand. 1996. Relationship between copper exposure duration, tissue copper concentration, and Rainbow Trout growth. Aquatic Toxicology 36:17–30.

McIntyre, J. K., D. H. Baldwin, J. P. Meador, and N. L. Scholz. 2008. Chemosensory deprivation in juvenile Coho Salmon exposed to dissolved copper under varying water chemistry conditions. En-vironmental Science and Technology 42:1352-1358.

McIntyre, J. K., D. H. Baldwin, D. A. Beauchamp, and N. L. Scholz. 2012. Low-level copper exposures increase visibility and vulner-ability of juvenile Coho Salmon to Cutthroat Trout predators. Ecological Applications 22:1460-1471.

McKenzie, L. M., R. Z. Witter, L. S. Newman, and J. L. Adgate. 2012. Human health risk assessment of air emissions from develop-ment of unconventional natural gas resources. Science of the Total Environment 424:79-87.

NRC (National Research Council). 2005. Superfund and mining megasites: lessons from the Coeur d’Alene River Basin. National Academies Press, Washington, DC.

Orr, J. C., V. J. Fabry, O. Aumont, L. Bopp, S. C. Doney, R. A. Feely, A. Gnanadesikan, N. Gruber, A. Ishida, F. Joos, R. M. Key K. Lindsey, E. Maier-Reimer, R. Matear, P. Monfrey, A. Mouchet, R. G. Naj-jar, G.-K. Plattner, K. B. Rodgers, C. L. Sabine, J. L. Sarmiento, R. Schiltzer, R. D. Slater, I. J. Totterdell, M.-F. Weirig, Y. Yamanaka, and A. Yoo. 2005. Anthropogenic ocean acidification over the twenty-first century and its impact on calcifying organisms. Na-ture 437:681-686.

Pond, G. J., M. E. Passmore, F. A. Borsuk, L. Reynolds, and C. J. Rose. 2008. Downstream effects of mountaintop coal mining: com-paring biological conditions using family- and genus-level mac-roinvertebrate bioassessment tools. Journal of the North Ameri-can Benthological Society 27:717–737.

Smith, D. R., C. D. Snyder, N. P. Hitt, J. A. Young, and S. P. Faulkner. 2012. Shale gas development and Brook Trout: scaling best management practices to anticipate cumulative effects. Envi-ronmental Practice 14:1-16.

Sprague, J., P. Elson, and R. Saunders. 1965. Sublethal copper–zinc pollution in a salmon river—a field and laboratory study. Interna-tional Journal of Air and Water Pollution 9:531–543.

Stacey, S. L., L. L. Brink, J. C. Larkin, Y. Sadovsky, B. D. Goldstein, B. R. Pitt, and E. O. Talbott. 2015. Perinatal outcomes and unconven-tional natural gas operations in Southwest Pennsylvania. PLOS One DOI: 10.1371/journal.pone.0126425.

USEPA (U.S. Environmental Protection Agency). 1995. Human health and environmental damages from mining and mineral process-ing wastes. Office of Solid Waste, Washington DC.

_____. 2000. Liquid assets: America’s water resources at a turning point. EPA-840, Washington, DC.

_____. 2011. The effects of mountaintop mines and valley fills on aquatic ecosystems of the central Appalachian coalfields. Office of Research and Development, EPA/600/R-09/138F, Washing-ton, D.C.

_____. 2014. An assessment of potential mining impacts on salmon ecosystems of Bristol Bay, Alaska. EPA 910-R-14-001A-C, Wash-ington, D.C.

Uechi, Jenny. 2014. Imperial Metals Mount Polley disaster could cost $500 million, but bonds only a fraction of this amount. Vancou-ver Observer (August 20). Available: www.vancouverobserver.com/news/imperial-metals-mount-polley-disaster-could-cost-500-million-bonds-only-fraction-amount. (August 2015).

Vox. 2015. How the EPA managed to spill 3 million gallons of mining waste into a Colorado River. Available: www.vox.com/2015/8/10/9126853/epa-mine-spill-animas. (August 2015).

Webb, E., S. Bushkin-Bedient, A. Chang, C. D. Kassotis, V. Balise, and S. C. Nagel. 2014. Developmental and reproductive effects of chemicals associated with unconventional oil and natural gas operations. Reviews on Environmental Health 29:307-318.

Weltman-Fahs, M., and J. M. Taylor. 2013. Hydraulic fracturing and Brook Trout habitat in the Marcellus Shale region: potential im-pacts and research needs. Fisheries 38(1):4-15.

WISE (World Information Service on Energy). 2011. Chronology of major tailings dam failures. WISE Uranium Project. Available: www.wise-uranium.org/mdaf.html. (August 2015).

Woody, C. A., R. M. Hughes, E. J. Wagner, T. P. Quinn, L. H. Roulsen, L. M. Martin, and K. Griswold. 2010. The U.S. General Mining Law of 1872: change is overdue. Fisheries 35(7):321-331.

• Call 800-843-1172 to discuss your custom tagging needs • Email us at [email protected] • View our website for our latest catalog www.floytag.com

The World Leader & Innovator in Fish Tags

floy tag ad3.indd 1 1/24/2013 6:45:34 PM

ESSAY


Defining Forage Species to Prevent a Management DilemmaKonstantine J. RountosDepartment of Biology, St. Joseph's College, 155 Roe Blvd., Patchogue, NY 11772. E-mail: [email protected]

MOTIVATION

Forage species are often defined in scientific and popular literature using terms such as “small,” “schooling,” “short-lived,” pelagic fish found at intermediate trophic levels of marine food chains. However, not all stakeholders use the same combination of terms, and their definitions include an array of fish and invertebrates that range in size, life span, and habitat preferences. Because forage species are ecologically important (e.g., Pikitch et al. 2012), often economically important (e.g., Pikitch et al. 2014), and increasingly promoted for direct human nutrition (e.g., Tacon and Metian 2013), it is both surprising and a concern that there is such an array of definitions being used. This diversity may cause undue confusion that will further complicate the sustainable management of these species.

The International Symposium on the Role of Forage Fishes in Marine Ecosystems held in Alaska determined that “forage fish is a concept that many people have come to understand because of the context it is used in, but for which we lack a concrete definition. The term embodies a peculiar combination of ambiguity and precision” (Springer and Speckman 1997:773). Nearly 20 years later, we still lack a common operational definition used among scientists, industry, policy makers, and the public. Finding a better definition is important, because there is not only a global interest in understanding the trade-offs and approaches needed to sustainably manage these species (e.g., Peck et al. 2014; Essington et al. 2015; Rountos et al. 2015), but also a need for identifying them for ecolabel certifications (e.g., Agnew et al. 2014). It is time that we created a consistent definition of forage species: what they are—and are not—to prevent a future dilemma.

APPROACH

This micro-analysis aimed to examine and analyze how different stakeholders have defined forage species in the past, in order to help identify and define those species in the future. Two approaches were used to compile and evaluate the various definitions of forage species being used. First, a literature search was conducted to explore the diversity of forage terminology and scientific criteria. Next, a search of the attributes (i.e., maximum total length, life expectancy, trophic level, habitat) of those species included in the literature search was carried out using FishBase (Froese and Pauly 2015) and other sources. The diversity of definitions in the results suggests that a standardized definition of forage species should (1) be more specific in life history attributes and (2) focus more on whether or not a species is providing a critical role as prey in marine ecosystems. The latter may be accomplished by establishing criteria based on dietary contributions of the forage species or using other trophodynamic indicators.

THE NAME GAME

There are many terms currently used to identify forage species, including “forage fish,” “forage species,” “small pelagics,” and “lower-trophic-level species.” Scientists who

use these terms often include a variety of fish and invertebrates (e.g., euphausiids, cephalopods, shrimp); thus, the term forage fish should immediately be phased out. The term that is most appropriate from a technical standpoint is forage species, because it is not exclusive to finfish, size, habitat, or trophic level.

TO INCLUDE OR NOT TO INCLUDE, THAT IS THE QUESTION

Scientists do not always agree on whether to include juvenile fish, myctophids, euphausiids (i.e., krill), cephalopods, or shrimp in their definitions of forage species. Most considerations have included krill, but there is not a clear consensus on other invertebrates such as cephalopods or shrimp. Some studies and management documents argue that cephalopods should not be included, because they can be quite piscivorous, whereas other studies show that certain species of cephalopods provide an important dietary component to upper-trophic-level predators (e.g., Szoboszlai et al. 2015) and thus should be categorized as forage species. Ultimately, this may depend on the species of cephalopod, because they represent a range of trophic levels (Coll et al. 2013). Inclusion of juvenile fish is also debatable, because they do not fulfill a role as prey throughout their entire life history. Juvenile Alaska Pollock Theragra chalcogramma, rockfish Sebastes spp., and salmon Oncorhynchus spp. are signature examples of fish that are important forage when juveniles but do not fit this criterion as adults. Myctophids, and possibly other small midwater fishes, are another group that are rarely included in forage species definitions, even though they are ecologically important as prey (e.g., Catul et al. 2011).

IS IT A “SMALL PELAGIC” WORLD AFTER ALL?

Categorization of species as forage has often depended on physical, ecological, and behavioral attributes. In particular, forage species are often considered small (<30 cm maximum standard length), relatively short-lived (1–3 years), pelagic fish that occupy intermediate trophic levels. However, there is ambiguity when the actual attributes of these species are cross-referenced using FishBase or other sources.

SizeMost studies that categorize forage species as small

pelagics (<30 cm) actually include species with maximum lengths ≥30 cm (e.g., Atlantic Menhaden Brevoortia tyrannus, etc.), according to FishBase. The median maximum length of forage species amongst the studies in the literature search is 28 cm, ranging from 2.5 to 76 cm. Therefore, more appropriate definitions should state that these species are small (<30 cm) to intermediate (≥30 but <90 cm) sized. Size-based classifications become more complicated if juvenile life stages of fish are included, because these species (e.g., rockfishes, mackerels, etc.) can have maximum lengths exceeding 90 cm. Because only 4% (i.e., 425 out of 9,992) of small (<30 cm) marine fish species in FishBase are from forage families (i.e., Atherinopsidae, Hemiramphidae, Dussumieriidae, Pristigasteridae, Osmeridae,

ESSAY


Clupeidae, Alestidae, Ammodytidae, Argentinidae, Centriscidae, Atherinidae, Stromateidae, and Engraulidae), using size in a definition may not be an ideal way to define forage species.

AgeSimilar discrepancies are found in regards to longevity.

Forage species are not as short-lived as many definitions have asserted, and this misconception is largely based on notable examples of short-lived forage species like the Peruvian Anchoveta Engraulis ringens (maximum age = 3). The median maximum age of forage species amongst the studies in this literature search was 10 years, ranging from 2 to 25. This point is further emphasized when juvenile fish are added as forage species, with many living a median maximum age of 18 years

HabitatPelagic is frequently used in definitions of forage species,

but there are examples where this attribute is not appropriate. Sandeels (family Ammodytidae), for instance, are not pelagic, yet they are a major forage species (Holland et al. 2005). Similarly, benthic invertebrates (e.g., polychaetes, amphipods, mysids, etc.) also serve as important forage in some ecosystems (Ihde et al. 2015). For fish, a search of forage families in FishBase revealed that nearly 81% (i.e., 457 out of 566) of forage species are pelagic, whereas 19% (i.e., 109 out of 566) are either reef-associated (11%) or demersal (8%). Although fish species are mostly pelagic, it is not an appropriate characteristic for defining all forage species.

Trophic LevelForage species are often defined as occupying intermediate

trophic levels, but some studies refer to them as lower-trophic-level species (e.g., Smith et al. 2011). Trophic levels of forage species included in this literature search ranged from 2.1 to 4.5, with a median trophic level of 3.2, which identifies as “intermediate.” The median trophic level increased to 3.6 when juvenile fish are included. These values would decrease if benthic invertebrates were included.

FINDING COMMON GROUND FOR AN OPERATIONAL DEFINITION

The goal of this literature search is not to single out or criticize the definitions used in specific studies but rather to emphasize the need to create a consistent operational definition for these species. Although the focus has been on the inconsistencies in species type, size, age, habitat, and trophic level, it is important to note that when defining forage species, every study indicated the ecological importance of these species to upper-trophic-level predators. Therefore, a common standard should focus more on the trophic role of a species and whether it is a critical prey resource throughout its life history. This could be done by implementing a dietary component into the definition of forage species or by using indicators such as the “SURF” index (Plagányi and Essington 2014). Diet studies and syntheses of diet data for upper-trophic-level predators can provide guidance to scientists, managers, and policy makers on which species are likely to be forage species. It is imperative that we are aware, critical, and consistent in how we define forage species in our future work, because definitions are paramount for communication, legislation, and effective ecosystem-based management (Link and Browman 2014).

ACKNOWLEDGMENTS

The editors and Ed Houde are thanked for their thoughtful review and comments on an earlier draft of this article. Thanks are also offered to those who visited and interacted with my poster on this topic at the 2015 AFS Annual Meeting in Portland, Oregon. Rachel Silver and Alexandra DiGiacomo are especially thanked for their assistance with the FishBase searches.

REFERENCESAgnew, D. J., N. L. Gutiérrez, A. Stern-Pirlot, and D. D. Hoggarth. 2014.

The MSC experience: developing an operational certification standard and a market incentive to improve fishery sustainability. ICES Journal of Marine Science 71(2):216–225.

Catul, V., M. Gauns, and P. K. Karuppasamy. 2011. A review on mes-opelagic fishes belonging to family Myctophidae. Reviews in Fish Biology and Fisheries 21(3):339–354.

Coll, M., J. Navarro, R. J. Olson, and V. Christensen. 2013. Assessing the trophic position and ecological role of squids in marine eco-systems by means of food-web models. Deep-Sea Research Part II-Topical Studies in Oceanography 95:21–36.

Essington, T. E., P. E. Moriarty, H. E. Froehlich, E. E. Hodgson, L. E. Koehn, K. L. Oken, M. C. Siple, and C. C. Stawitz. 2015. Fishing amplifies forage fish population collapses. Proceedings of the National Academy of Sciences 112(21):6648–6652.

Froese, R., and D. Pauly. 2015. FishBase. Available: www.fishbase.org. (July 2015).

Holland, G. J., S. P. Greenstreet, I. M. Gibb, H. M. Fraser, and M. R. Rob-ertson. 2005. Identifying sandeel Ammodytes marinus sediment habitat preferences in the marine environment. Marine Ecology Progress Series 303:269–282.

Ihde, T. F., E. D. Houde, C. F. Bonzek, and E. Franke. 2015. Assessing the Chesapeake Bay forage base: existing data and research pri-orities. STAC Publication Number 15-005, Chesapeake Bay Pro-gram, Edgewater, Maryland.

Link, J. S., and H. I. Browman. 2014. Integrating what? Levels of marine ecosystem-based assessment and management. ICES Journal of Marine Science 71(5): 1170–1173. DOI: 10.1093/icesjms/fsu026.

Peck, M. A., S. Neuenfeldt, T. E. Essington, V. M. Trenkel, A. Takasuka, H. Gislason, M. Dickey-Collas, K. H. Andersen, L. Ravn-Jonsen, N. Vestergaard, S. F. Kvamsdal, A. Gårdmark, J. Link, and J. C. Rice. 2014. Forage fish interactions: a symposium on “creating the tools for ecosystem-based management of marine resources.” ICES Journal of Marine Science 71(1):1–4.

Pikitch, E. K., P. D. Boersma, I. L. Boyd, D. O. Conover, P. Cury, T. E. Ess-ington, S. S. Heppell, E. D. Houde, M. Mangel, D. Pauly, É. Plagányi, K. Sainsbury, and R. Steneck. 2012. Little fish, big impact: manag-ing a crucial link in ocean food webs. Lenfest Ocean Program, Washington, D.C.

Pikitch, E. K., K. J. Rountos, T. E. Essington, C. Santora, D. Pauly, R. Watson, U. Sumaila, P. D. Boersma, I. L. Boyd, D. O. Conover, P. Cury, S. S. Heppell, E. D. Houde, M. Mangel, É. Plagányi, K. Sains-bury, R. Steneck, T. M. Geers, N. Gownaris, and S. B. Munch. 2014. The global contribution of forage fish to marine fisheries and ecosystems. Fish and Fisheries 15:43–64.

Plagányi, É. E., and T. E. Essington. 2014. When the SURFs up, forage fish are key. Fisheries Research 159:68–74.

Rountos, K. J., M. G. Frisk, and E. K. Pikitch. 2015. Are we catching what they eat? Moving beyond trends in the mean trophic level of catch. Fisheries 40(8):376–385.

Smith, A. D. M., C. J. Brown, C. M. Bulman, E. A. Fulton, P. Johnson, I. C. Kaplan, H. Lozano-Montes, S. Mackinson, M. Marzloff, L. J. Shan-non, Y.-J. Shin, and J. Tam. 2011. Impacts of fishing low-trophic level species on marine ecosystems. Science 333(6046):1147–1150.

Springer, A. M., and S. G. Speckman. 1997. A forage fish is what? Sum-mary of the symposium. Pages 773–816 in B. Baxter and C. W. Mecklenburg, editors. Forage fishes in marine ecosystems. Pro-ceedings of the international symposium on the role of forage fishes in marine ecosystems. University of Alaska, Alaska Sea Grant College Program Report Number 97-01, Fairbanks, Alaska.

Szoboszlai, A. I., J. A. Thayer, S. A. Wood, W. J. Sydeman, and L. E. Koehn. 2015. Forage species in predator diets: synthesis of data from the California Current. Ecological Informatics 29(1):45–56.

Tacon, A. G. J., and M. Metian. 2013. Fish matters: importance of aquatic foods in human nutrition and global food supply. Re-views in Fisheries Science 21(1):22–38.


FEATURE

Ray HilbornSchool of Aquatic and Fishery Sciences, Box 355020, University of Washington, Seattle, WA 98195. E-mail: [email protected]

Correlation and Causation in Fisheries and Watershed Management


INTRODUCTION

When natural resources, such as a fishery or an ecosystem, are managed in order to achieve a desired societal goal, it is necessary to change or manipulate certain elements of the resource. Managers change laws, regulations, expenditures, and actions, and each one of these changes will have a distinct effect. Fisheries and watershed management is no different from any other form of management, whether management of a national economy, a manufacturing plant, or a patient in a hospital. To predict the likely outcome of different actions, managers rely on a mixture of historical experience, theory, and intuition. Of these three elements, historical experience is considered the most reliable because the more often alternative actions have been tested in multiple applications, the more confident managers will be in the likely outcomes.

Historical experience can take two basic forms: observational and manipulative. Observational studies are far more common and summarize the historical experience of the relationship between two variables of interest. A classic example is the analysis of spawning stock and subsequent recruitment. Given a history of the size of a spawning stock and the subsequent recruitment, we can estimate how any manipulation of the size of a spawning stock will affect subsequent recruitment. In a summary of historical knowledge of the relationship between spawning stock and recruitment, Myers et al. (1994) concluded that there was strong evidence

Efforts to understand how to manage aquatic ecosystems often rely on correlations between human actions and impacts in the ecosystem. We are often warned that correlation does not imply causation and that the gold standard for identify-ing cause and effect relationships is manipulative experiments. History shows us that correlations are often not causal and that managers should not design policies based on the assumption of causality. However, in the absence of manipulation, correlative evidence may be all that is available. Correlative evidence is strongest when (1) correlation is high, (2) it is found consistently across multiple situations, (3) there are not competing explanations, and (4) the correlation is consist-ent with mechanistic explanations that can be supported by experimental evidence. Where possible, manipulative experi-ments and formal adaptive management should be employed, but in large-scale aquatic ecosystems these opportunities are limited. More commonly, we should emphasize identifying the range of possible causal mechanisms and identify poli-cies that are robust to the alternative mechanisms.

Correlación y causalidad en pesquerías y manejo de cuencas hidrográficasLos esfuerzos que se realizan para administrar los ecosistemas acuáticos, a veces se basan en correlaciones entre las acciones humanas y los impactos de éstas sobre los ecosistemas. Suele insistirse en que la correlación no implica cau-salidad y que el estándar crítico para identificar relaciones causa-efecto son los experimentos controlados. La historia muestra que las correlaciones no siempre son causales y que los manejadores no debieran diseñar políticas basadas en la suposición de existencia de causalidad. No obstante, en ausencia de manipulación, la evidencia correlativa puede ser lo único que hay disponible. La evidencia correlativa es más fuerte cuando (1) la correlación es alta, (2) se obtiene de forma consistente a través de múltiples situaciones, (3) no existen explicaciones alternativas, y (4) la correlación es consistente con explicaciones mecánicas que pueden ser apoyadas con evidencia experimental. Cuando sea posible, se deben utilizar los experimentos controlados y el manejo adaptativo formal, sin embargo en el caso de ecosistemas acuáticos de gran escala, estas oportunidades son limitadas. Más comúnmente, se debe hacer énfasis en identificar un rango de posibles mecanismos causales e implementar políticas de manejo que sean robustas a los mecanismos alternativos.

Corrélation et Causalité entre la Pêche et la Gestion des Bassins-versantsLes efforts pour comprendre comment gérer les écosystèmes aquatiques se basent souvent sur les corrélations entre les actions humaines et les impacts sur l'écosystème. Nous sommes souvent mis en garde sur le fait que la corrélation n’implique pas la causalité et que le critère de référence pour identifier des liens de cause à effet sont les expériences demanipulation. L’histoire nous enseigne que les corrélations sont souvent sans causalité et que les responsables ne devraient pas concevoir les politiques en se basant sur la présomption de causalité. Néanmoins, en absence de manipula-tion, la preuve de la corrélation est peut-être la seule chose disponible. La preuve de la corrélation est plus forte quand (1) la corrélation est importante, (2) elle est retrouvée systématiquement à travers de nombreuses situations, (3) il n’y a pas d’explications antagonistes, et (4) la corrélation est cohérente avec des explications mécanistes qui peuvent s’appuyer sur la preuve expérimentale. Si possible, des expériences de manipulation et la gestion formelle adaptative devraient être uti-lisées, mais dans les écosystèmes aquatiques à grande échelle ces opportunités sont limitées. Généralement, nous devons mettre l’accent sur l’identification de l’habitat des possibles mécanismes de causalité et identifier les politiques qui sont robustes pour les mécanismes alternatifs.

that reducing spawning stock size below certain levels will lead to reduced recruitment. This relationship between low spawning stock size and low recruitments was observational evidence that supported the move in the United States and other countries to prevent low spawning stock size and increase the abundance of stocks that were at low density.

In this case, the correlation between low spawning stock size and low recruitment has generally been interpreted as being causal, and throughout much of fisheries and watershed management we often assume that correlation means causation. Yet it is also widely accepted that correlation does not necessarily mean causation. Should we then find the strongest correlations and assume causation?

Learning from manipulation is the gold standard in learning from experience, and in fisheries management much has been learned from manipulation. Hatchery managers deliberately experiment with time or size of release, feeds, and rearing densities. Freshwater fisheries managers experiment with stocking times and densities. In both cases, these manipulations are often controlled; that is, alongside the experiment there is another population that is thought to be largely similar but not given the treatment of interest. Any manipulative experiments should ideally have three elements—controls, replication, and randomization of treatments—and the more units available to carry out experiments, the more likely these three elements will be achieved.


As we move to larger spatial scales, particularly in marine fisheries and watershed management, the opportunities for controls and replication are fewer. There are hundreds of marine fisheries and watersheds around the United States, and though they may contain the same species, each one is geographically and environmentally different enough to weaken any comparison or potential for manipulation. Even large-scale meta-analyses involving dozens or hundreds of data sets as in Myers et al. (1994) have their flaws. In response to the Myers et al. (1994) argument that low spawning stocks lead to low recruitment, Gilbert (1997) argued that the causal relationship lies elsewhere. Poor environmental conditions actually cause sustained low recruitment that leads to lower spawning stocks, and increasing spawning stock size by dramatically reducing fishing pressure will not result in higher recruitment. This has recently received support from Szuwalski et al. (2015), who analyzed over 200 data sets and found that environmental regime changes provide a better explanation for changes in recruitment than changes in spawning stock size.

In many cases, fishery managers have only the experience in a single fishery or watershed of interest to guide them as they seek to determine what factors lead to better outcomes. Examining the historical evidence will often point toward a correlation between environmental conditions and the productivity of a fish stock. Then we must ask, if such an environmental condition could be affected by management, will manipulating it lead to better stock size or production?

This is a question of causality. Does the environmental condition really cause a change in stock productivity? Though everyone is familiar with “correlation does not imply causation,” correlative studies have often been the foundation for management actions in fisheries and other fields and, in the absence of other information, managers are certainly tempted to assume that correlation at least suggests causation.

The nature of causation is one of the oldest questions in philosophy, and though much of the theory is either abstract or highly mathematical, it is quite relevant to fisheries. In this article, we will review the theory of causation, examine the relationship between correlation and causation, and evaluate the current state of how to approach fisheries management when the major evidence is correlative. To do this, I will review some examples from outside fisheries where correlation proved to also be causation (cigarette smoking and cancer) and where correlation was not causation (hormone replacement therapy and heart disease) and relate the lessons from these studies to fisheries and watershed management.

THEORY

Philosophical discussions of the nature of causation date back at least to Aristotle, who distinguished between different kinds of causes (Holland 1986). Locke (1690; cited in Holland 1986:950) proposed: “That which produces any simple or complex idea, we denote by the general name ‘cause’, and that which is produced, ‘effect.’” This philosophy seems to provide little guidance for managers who must make decisions. Of much more practical use is the statistical literature. Rubin (1974) suggested that randomized trials are by far the most powerful form of evidence and, despite opposition from many of his contemporaries, argued that nonrandomized trials can also be informative. A most important point in using any historical experience is that “trials in the study are representative of other future trials” (Rubin 1974:697). There are two key parts to this

assertion: one is often called stationarity, in which relationships from the past will be the same in the future. The second is that the manipulated experimental units, such as fish stocks or watersheds, represent the kind of units that will be managed in the future. However, Rubin (1974) also pointed out that observational studies have advantages over randomized trials because they generally observe systems in a broader set of states than randomized experiments. Experimental units should be as similar as possible, thus representing only a small fraction of conditions in the real world. Finally, Rubin (1974) argued that nonrandomized studies can be very powerful if the analyst can successfully assert that there is only one variable—the one of interest—that may causally affect the outcome.

Hill (1965) evaluated the relationship between correlation and causation in public health and medicine and identified three primary characteristics that provided increased evidence for a causal correlation. The first is strength of association: the higher the correlation, the greater the belief that the factor is causal. The second is consistency: when looking across populations or units of observation, the more the factor is consistently associated, the greater the belief. Finally, there is specificity, which, in the biomedical world, is a lack of other correlations or plausible explanations. The same is true in fisheries. If any other variables also correlate with the effect of interest, there is less specificity and thus weaker evidence for causation.

Holland (1986) developed a model of causal inference almost exclusively for randomized trials, but he also reviewed the literature on nonrandomized trials and observational studies and emphasized that we should look more for the effects of causes, rather than the causes of effects, which fits nicely into the framework of management and decision theory. We want to know what will happen if we manipulate the system (change the causes) more than we want to know what causes the effect.

As we know more about the biology of a system, the actual cause may be refined. For example, do bacteria cause disease? Well, yes … until we dig deeper and find that it is the toxins the bacteria produce that really cause the disease; and this is really not it either. Certain chemical reactions are the real causes … and so on, ad infinitum. (Holland 1986:959)

What the manager wants to know is, if he prevents contact with the bacteria, will there be no disease—not the actual cause of the disease.

The essence of Rubin and Holland for fisheries and watershed managers is summarized in the phrase, “No causation without manipulation” (Holland 1986:959).

Though everyone is familiar with “correlation does not imply causation,” correlative stud-ies have often been the foundation for man-agement actions in fisheries and other fields and, in the absence of other information, managers are certainly tempted to assume that correlation at least suggests causation.


CLASSIC EXAMPLES OF CORRELATION

Two examples that highlight the potential and perils of using correlation for public policy are found in the medical literature. That smoking causes lung cancer is now widely accepted, whereas that hormone replacement therapy reduces heart disease turned out to be a false correlation. Worse yet, randomized clinical trials showed that hormone replacement therapy actually increased heart disease.

Smoking and Lung CancerDuring the first half of the 20th century, the incidences

of lung cancer increased steadily and various explanations were put forward (White 1990), including cigarette smoking, industrial pollution, smoke from domestic fires, and tars used in road construction. The least persuasive evidence was that the frequency of smoking increased at the same time that lung cancer increased. Stronger evidence was that smokers were much more likely to develop lung cancer than nonsmokers (Doll and Hill 1950). But the strength of the evidence was much criticized. Most interesting, from a historical perspective, were the many papers by Sir R. A. Fisher, perhaps the best known statistician of the 20th century. Fisher (a heavy smoker) argued quite correctly that there were two reasons why the correlation may not be causal.

Two classes of alternative theories which any statistical association, observed without the precautions of a definite experiment, always allows—namely, (1) that the supposed effect is really the cause, or in this case, that incipient cancer, or a pre-cancerous condition with chronic inflammation, is a factor in inducing the smoking of cigarettes, or (2) that cigarette-smoking and lung cancer, though not mutually causative, are both influenced by a common cause, in this case the individual genotype. (Fisher 1957:297).

However, throughout the 1950s and 1960s, evidence continued to increase, including data showing that the incidence of lung cancer was indeed related to the frequency of smoking, that those who quit smoking showed a reduced incidence of lung cancer compared to those who continued, and, finally, the demonstration in laboratory studies that elements of tobacco smoke could induce cancer in laboratory animals. By 1959, the U.S. Surgeon General concluded that “(t)he weight of evidence at present implicates smoking as the principal etiological factor in the increased incidence of lung cancer” (Burney 1959:141). Yet the matter was not altogether laid to rest. In an editorial, the editor of the Journal of the American Medical Association said, “A number of authorities … do not agree with his conclusion” (Talbot 1959:162), but by 1964, both the Surgeon General and the Journal of the American Medical Association had made public statements that smoking was a health hazard.

Throughout the entire discussions of the 1950s, it was always understood that, unless a causal mechanism was identified, any correlation would be questionable. “The definitive investigations,” Berkson (1960:369) wrote, “must come from the biologic sciences, pathology, pharmacology, chemistry, and so forth. … We will not really know whether smoking causes cancer, till we know at least something in a precise way about how it causes cancer.” Throughout the 1960s and 1970s, the carcinogenic effects of elements of tobacco smoke were more and more demonstrated in vitro and in vivo

(Miller and Miller 1981). Of course, the definitive evidence would have had to come from randomly assigning individual humans to smoking and nonsmoking groups, an ethically indefensible experiment. In experiments with laboratory animals, though, the evidence is strong.

Hormone Replacement TherapyIn his 1966 bestseller Feminine Forever, Robert Wilson

argued that menopause was a treatable phenomenon and women could take hormones to replace the ones their bodies were no longer producing. Most women would certainly love to avoid consequences of menopause such as hot flushes, weight gain, decreased sexual desire, and masculinization, and when Wilson told them that this could all be avoided with simple hormone replacement therapy (HRT), the news was greeted with great joy. Then the news got even better. Comparisons of women who took HRT with those who did not indicated that HRT reduced heart disease and hip fractures (Grady et al. 1992). Though this evidence was correlative, the lure of a technical solution to an undesirable condition was such that HRT was widely prescribed and used in developed countries.

However, unlike with smoking, randomized trials with HRT were possible and were conducted in the 1980s. Women who sought treatment were randomly assigned to either HRT or a placebo in double-blind trials, and their medical history was followed for a number of years. The not-so-joyous results were published in 1998 (Hulley et al. 1998): the frequency of heart disease was in fact higher in the HRT treatment group, as was the risk of breast cancer, stroke, and blood clots. Moreover, the women on HRT in the earlier, correlative studies had been richer and better educated, had better diets, exercised more, and had better access to medical care than those not on HRT (Lawlor et al. 2004a, 2004b), which accounted for their general better health and counteracted the rather modest negative effects of HRT. The HRT did not reduce heart disease, but it appeared to do so in correlative analysis because those who received HRT were healthier than those who did not receive HRT. Correlation was not causation, but the only way it was identified was by manipulation.

Fish Examples One holy grail for fisheries scientists has been understanding

the fluctuations in the recruitment of fish stocks. We know that survival of eggs and larvae through the first year of life depends greatly on the availability of food, shelter from predators, and for many species favorable currents to either transport larvae to the appropriate habitat or to retain them where they were born. Throughout the 20th century, fisheries scientists looked for measureable environmental conditions that correlate with recruitment success by searching data sets of environmental variables. Shepherd et al. (1984) reviewed 47 studies that had found correlations between environmental variables and recruitment success. Myers (1998:298) looked at each of these stocks again and found that by the mid-1990s only one of the 47 relationships was used by assessment scientists because the correlations that existed in the data up to 1984 had failed to reliably predict recruitment in later years. With more data, the correlations fell apart. The exceptions generally were stocks at the limit of their geographic range, where some correlations with temperature were consistent. Myers concluded that, on the whole, using environmental correlates should be avoided when choosing management actions: “The rarity of the use of environment/recruitment correlations is clear evidence against their general usefulness in assessments. Even if an


environmental variable is important, it does not mean that it is key to the management of the fishery.”

A different way of looking at what determines fish recruitment has been analysis of the relationship (i.e., correlation) between spawning stock and recruitment. Ricker’s classic 1954 “Stock and Recruitment” is one of the most cited papers in the fisheries literature, and for many agencies the spawner–recruit relationship is one of the foundations for setting management policies. Though conceptually one can think of a spawning stock as being manipulated, in fact, almost all stock recruitment analyses look for a repeatable relationship (again, a correlation) between number of spawners and number of recruits. Myers et al. (1994) presented a meta-analysis of spawner–recruit data of 72 finfish stocks. They concluded that there was indeed evidence that lower spawning stocks produced lower recruitment, and one could identify spawning stock thresholds below which lower recruitment could be expected. They said that their analysis “should help dispel the widely held notion that observed recruitment is usually independent of spawning biomass. … The mean recruitment is generally greater above the threshold than below” (Myers et al. 1994:203). Their argument was that higher spawning stock size on average was associated with higher recruitment; therefore, higher spawning stock size caused higher recruitment. This would be a logical assumption. However, should the opposite be true, rather strong density-dependent processes must be operating.

On the other hand, we do know that there are lots of mechanisms leading to strong density dependence, and Gilbert (1997) proposed an alternative hypothesis. He suggested that recruitment is largely driven by periodic shifts in environmental factors and that recruitment will be high when conditions are good and poor when conditions are bad. A sustained period of good recruitment will result in a sequence of high recruitments, which in turn leads to high spawning stock. Conversely, a period of poor environmental conditions will result in a series of poor recruitments that will lead to low spawning stocks. Thus, Gilbert (1997) proposed that low recruitment causes low spawning stock and high recruitment causes high spawning stock and that when spawning stocks are low, increasing them is unlikely to increase recruitment.

This interpretation has not been generally accepted in fisheries management agencies, although a greater number of assessments now include periodic shifts in stock productivity when calculating management reference points. However, in support of Gilbert’s perspective, Szuwalski et al. (2015) looked at several hundred data sets both of spawner–recruit relationships and biomass-to-surplus-production relationships and concluded that the behavior of far more stocks is consistent with period regimes of good and bad conditions rather than biomass leading to stock productivity.

There have been at least two deliberate manipulative experiments in Pacific salmon. Based on analysis of spawner–recruit data, the number of Sockeye Salmon Oncorhynchus nerka allowed to spawn in Rivers Inlet, British Columbia (Walters et al. 1993), and in the Kvichak River, Alaska (Eggers and Rogers 1987), were increased. In both cases, recruitment did not increase and the manipulative experiments failed to demonstrate causality.

Moving from single species to ecosystems, there is increasing use of mechanistic models as a way to evaluate ecosystem impacts—Ecopath with Ecosim (Christensen and Walters 2004) and ATLANTIS (Fulton et al. 2005). These models do not rely on correlation in their construction but do

rely on correlation of their predicted values to observed data as informal validation of the mechanisms. Thus, though they represent highly complex hypotheses, they have the same characteristics of simple (A causes B) hypotheses with respect to validation of predicted impacts of management policies. When correlative evidence is weak, manipulation is the strongest test.

Some of the counterintuitive results can be explained by the work of George Sugihara and his students, who emphasized the nonlinearity of ecological systems (Glaser et al. 2011; Sugihara et al. 2011) and used simple nonlinear models to demonstrate both that lack of correlation does not mean lack of causation and that correlation does not mean causation. In these models, it is easily demonstrated that, although species A feeds on species B and fishing down species A should result in an increase in the abundance of species B, observation of time series data will often show no correlation between A and B. In summary, the fisheries literature suggests that in general correlation has not meant causation.

Management ApproachesIf we take “no causation without manipulation” literally,

most fisheries management reaches a dead end because almost all management systems have enough unique features that make replication and control impossible. However, the theory of adaptive management (Walters 1986) can provide guidance on how to proceed. Only when there is the potential for spatial or temporal replication can true experimental designs be established and experiments performed. But these cases are unusual enough that adaptive management that involves noncontrolled experiments may provide the best approach. Right away, a fishery manager has the key advantage of little concern with any of the details of causation. What matters are the effects of manipulation. A manager wants to know whether a system will respond differently to implementation or nonimplementation of a management action.

Adaptive management should begin with modeling that “is intended to serve three functions: (1) problem clarification and enhanced communication among scientists, managers, and other stakeholders; (2) policy screening to eliminate options that are most likely incapable of doing much good, because of inadequate scale or type of impact; and (3) identification of key knowledge gaps that make model predictions suspect” (Walters 1997:2). The key result of the modeling step is identification of alternative hypotheses about the dynamics of the system and an evaluation of the degree of support for each hypothesis. Guided by Hill (1965), we recognize that the more competing explanations there are, the less support correlation provides for causation. The second step in adaptive management is the design of management manipulations. With the help of modeling, the expected value in terms of both costs and benefits of alternative actions can be calculated for any set of management actions. This is a form of risk analysis, and the product should be an experimental design. The third step is implementation, and the fourth is evaluation. Most well-designed adaptive management plans will use information gained early in the process to guide later management actions.

It is in the modeling and design stages that the correlation issue will need to be confronted. If alternative models that allow for noncausation are not considered, correlation will likely be interpreted as causation. For instance, in the original work on fisheries adaptive management, Walters and Hilborn (1976) explored the uncertainty in spawner–recruit relationships but never considered Gilbert’s hypotheses about changing


recruitment causing changed spawning stock. Thus, they never questioned that there was a relationship between spawning stock size and recruitment; they only tried to reduce the uncertainty in what that relationship was. In the light of what we know now, a similar analysis would consider a much broader range of hypotheses.

A widely praised adaptive management program aims to maintain and restore valuable resources in the riparian zone of the Colorado River in the Grand Canyon (Meretsky et al. 2000; Walters et al. 2000). One experimental treatment was “mechanical removal” (i.e., electrofishing) of Rainbow Trout O. mykiss and Brown Trout Salmo trutta because they are known to eat juveniles of the endangered Humpback Chub Gila cypha. A major removal treatment started in 2004 and was followed by substantial increases in chub recruitment. However, in the same year, the Colorado River warmed up considerably due to low reservoir stages at the head of the canyon in Lake Powell, which led to releases of water from in and above the reservoir thermocline. Such warmer water was also expected to benefit chub by reducing vulnerability to predation and increasing juvenile growth rates to sizes where predation is lower.

Warm reservoir releases and low trout abundances persisted until 2012 and only in 2013 did the system start returning to a high trout–cold water state. Only now, when cold water conditions are likely to last for a few years, is it possible to replicate the initial trout treatment to see whether trout reduction alone is sufficient to insure good Humpback Chub recruitment. This is a great example of why replication, even though it can be very slow to achieve, is critical in large-scale experiments. This kind of scenario is likely to be typical of ecological systems in general.

Thus, though adaptive management does hold promise for resolving uncertainty, beware of its many pitfalls. Most of the problems are rooted in the design of replicated experiments, and we have to admit that for large-scale fisheries and watersheds the difficulties of randomization, replication, and controls are often just too overwhelming. The net result is that we will almost always make decisions faced with great uncertainty and have to recognize the range of possible causal mechanisms.

SUMMARY

Fisheries and watershed management is decision making under uncertainty, and when actions are proposed, their consequences should be evaluated across as many possible hypotheses as possible. The weight of evidence of each causal pathway needs to be assessed, and ideally, historical data on randomized manipulative experiments will be the strongest form of evidence. Unfortunately, we will often only have observational evidence, and the evaluation of the support for alternative causal pathways will almost always be subjective. Though strong correlation is good evidence (Hill’s strength of association), it should not be the primary factor in determining strength of evidence for any causal pathway. Equally important are consistency (when factor X changes, output Y changes consistently) and specificity (are there alternative hypotheses?). The plausibility of mechanisms adds support for any causal pathway.

Plowright et al. (2008:424) warned that “in practice, looking for positive evidence for a favored theory can be a useful first strategy, but can harden into a bias that prevents the evaluation of alternate explanations for patterns that surface as more data become available” and emphasized that one should adopt

the method of multiple working hypotheses and evaluate the evidence for each of the hypotheses simultaneously—a method also recommended in Hilborn and Mangel (1997). Perhaps the greatest error would be to find the strongest correlation, assume that it is causal, and ignore other possible causal mechanism.

The most useful conclusion from the range of studies in ecology and other fields is that maintaining multiple working hypotheses throughout the analysis is the most important guidance, that managers should identify policies that are robust to the range of alternative hypotheses (Schindler and Hilborn 2015), and that correlative evidence should be regarded a priori as weak support for causation.

ACKNOWLEDGMENT

I thank Carl Walters for his perspective on this issue and the specific adaptive management examples he provided.

REFERENCES Berkson, J. 1960. Smoking and cancer of the lung. Proceedings of

the Staff Meetings of the Mayo Clinic 35:367–385.Burney, L. E. 1959. Smoking and lung cancer—a statement of the

Public-Health Service. Journal of the American Medical Associa-tion 171:1829–1837.

Christensen, V., and C. Walters. 2004. Ecopath with Ecosim: methods capabilities and limitations. Ecological Modeling 172:109–139.

Doll, R., and A. B. Hill. 1950. Smoking and carcinoma of the lung—preliminary report. British Medical Journal 2:739–748.

Eggers, D. M., and D. E. Rogers. 1987. The cycle of runs of Sockeye Salmon (Oncorhynchus nerka) to the Kvichak River, Bristol Bay, Alaska: cyclic dominance or depensatory fishing? Canadian Spe-cial Publication in Fisheries and Aquatic Sciences 96:343–366.

Fisher, R. A. 1957. Dangers of cigarette-smoking. British Medical Journal 2:297–298.

Fulton, E. A., A. D. M. Smith, and A. E. Punt. 2005. Which ecological indicators can robustly detect effects of fishing? ICES Journal of Marine Science 62:540–551.

Gilbert, D. J. 1997. Towards a new recruitment paradigm for fish stocks. Canadian Journal of Fisheries and Aquatic Sciences 54:969–977.

Glaser, S. M., H. Ye, M. Maunder, A. MacCall, M. Fogarty, and G. Sugi-hara. 2011. Detecting and forecasting complex nonlinear dynam-ics in spatially structured catch-per-unit-effort time series for North Pacific Albacore (Thunnus alalunga). Canadian Journal of Fisheries and Aquatic Sciences 68:400–412.

Grady, D., S. M. Rubin, D. B. Petitti, C. S. Fox, D. Black, B. Ettinger, V. L. Ernster, and S. R. Cummings. 1992. Hormone-therapy to prevent disease and prolong life in postmenopausal women. Annals of Internal Medicine 117:1016–1037.

Hilborn, R., and M. Mangel. 1997. The ecological detective: confront-ing models with data. Princeton University Press, Princeton, New Jersey.

Hill, A. B. 1965. The environment and disease: association or causa-tion? Proceedings of the Royal Society of Medicine 58:295–300.

Holland, P. W. 1986. Statistics and causal inference. Journal of the American Statistical Association 81:945–960.

Hulley, S., D. Grady, T. Bush, C. Furberg, D. Herrington, B. Riggs, E. Vittinghoff. 1998. Randomized trial of estrogen plus progestin for secondary prevention of coronary heart disease in postmen-opausal women. Journal of the American Medical Association 280:605–613.

Lawlor, D. A., G. D. Smith, and S. Ebrahim. 2004a. Commentary: the hormone replacement–coronary heart disease conundrum: is this the death of observational epidemiology? International Journal of Epidemiology 33:464–467.

–––––. 2004b. Socioeconomic position and hormone replacement therapy use: explaining the discrepancy in evidence from obser-vational and randomized controlled trials. American Journal of Public Health 94:2149–2154.

Meretsky, V. J., D. L. Wegner, and L. E. Stevens. 2000. Balancing endangered species and ecosystems: a case study of adaptive management in Grand Canyon. Environmental Management 25:579–586.

Miller, E. C., and J. A. Miller. 1981. Searches for ultimate chemical car-cinogens and their reactions with cellular macromolecules. Can-cer 47:2327–2345.


Myers, R. A. 1998. When do environment–recruitment correlations work? Reviews in Fish Biology and Fisheries 8:285–305.

Myers, R. A., A. A. Rosenberg, P. M. Mace, N. Barrowman, and V. R. Restrepo. 1994. In search of thresholds for recruitment overfish-ing. ICES Journal of Marine Science 51:191–205.

Plowright, R. K., S. H. Sokolow, M. E. Gorman, P. Daszak, and J. E. Foley. 2008. Causal inference in disease ecology: investigating ecological drivers of disease emergence. Frontiers in Ecology and the Environment 6:420–429.

Ricker, W. E. 1954. Stock and recruitment. Journal of the Fisheries Research Board of Canada 11:559–623.

Rubin, D. B. 1974. Estimating causal effects of treatments in rand-omized and nonrandomized studies. Journal of Educational Psy-chology 66:688–701.

Schindler, D. E., and R. Hilborn. 2015. Prediction, precaution, and policy under global change. Science 347:953–954.

Shepherd, J. G., J. G. Pope, and R. D. Cousens. 1984. Variations in fish stocks and hypotheses concerning their links with climates. Rapports et Proces-Verbaux des Reunions du Conseil Interna-tional pour l’Exploration de la Mer 185:255–267.

Sugihara, G., J. Beddington, C. H. Hsieh, E. Deyle, M. Fogarty, S. M. Glaser, R. Hewitt, A. Hollowed, R. M. May, S. B. Munch, C. Per-retti, A. A. Rosenberg, S. Sandin, and H. Ye. 2011. Are exploited fish populations stable? Proceedings of the National Academy of Sciences 108:E1224–E1225.

Szuwalski, C. S., K. A. Vert-pre, A. E. Punt, T. A. Branch, and R. Hil-born. 2015. Examining common assumptions about recruit-ment: a meta-analysis of recruitment dynamics for worldwide marine fisheries. Fish and Fisheries 16(4): 633-648. DOI: 10.1111/faf.120831.

Talbot, J. 1959. Smoking and lung cancer. Journal of the American Medical Association 171:2104.

Walters, C. J. 1986. Adaptive management of renewable resources. Macmillian Publishing Co., New York.

Walters, C. J. 1997. Challenges in adaptive management of riparian and coastal ecosystems. Conservation Ecology [online serial] 1(2). DOI:10.1016/j.ecolecon.2015.04.027

Walters, C. J., R. D. Goruk, and D. Radford. 1993. Rivers Inlet sockeye salmon: an experiment in adaptive management. North Ameri-can Journal of Fisheries Management 13(2):253–262.

Walters, C. J., and R. Hilborn. 1976. Adaptive control of fishing sys-tems. Journal of the Fisheries Research Board of Canada 33:145–159.

Walters, C. J., J. Korman, L. E. Stevens, and B. Gold. 2000. Ecosystem modeling for evaluation of adaptive management policies in the Grand Canyon. Conservation Ecology [online serial] 4(2):1. www.consecol.org/vol4/iss2/art1/

White, C. 1990. Research on smoking and lung-cancer—a landmark in the history of chronic disease epidemiology. Yale Journal of Biology and Medicine 63:29–46.

Wilson, R. A. 1966. Feminine forever. M. Evans, New York.

2016 AFS International Fisheries Section Fellow Award Applications Due March 6, 2016

Attention Students and Young Professionals:The International Fisheries Section (IFS) of the American Fisheries Society (AFS) is excited to announce the 2016 International Fisheries Section Fellow Award. The Fellow will represent the International Fisheries Section of AFS by presenting a poster or oral

presentation at the 2016 Fisheries Society of the British Isles (FSBI) annual symposium, July 18-22, Bangor University, United Kingdom. The theme of the 2016 FSBI annual symposium is “Fish, Genes, and Genomes: Contributions to Ecology, Evolution, and Management.” For more information go to: fsbi.org.uk/conference-2016/symposium-theme-2.

Benefits:• FSBI presentation (a presentation slot has been

reserved for the recipient of this award)• Up to $2,100 (USD) of reimbursable travel expenses

associated with attending the FSBI annual symposium• Meeting registration, dormitory accommodations, and

meal stipend while at the symposium• 1 year of membership to FSBI• 2 years of leadership in the IFS

Eligibility:This award is open to all student and young professional members (within three years of graduation) of the AFS, especially those working on topics of international interest or those associated with the symposium theme. However, we encourage all interested student and young professional members to apply. Current membership with AFS is mandatory to apply for and receive this award.

Application:• To apply, please submit the following information as

a single PDF document to the committee chair of the IFS Fellow Program, Zach Penney ([email protected]), and the deputy committee chair, Jeremy Higgs ([email protected]), by March 6, 2016:

• Letter of interest (please include information about your involvement in AFS)

• Proposed FSBI presentation title and abstract• Concise resume• Proposed travel budget (i.e. airfare, ground

transportation, etc.); please specify availability of supporting funds you may have from other sources if budget exceeds allowance

Obligations:The awarded Fellow is expected to fulfill the following obligations:• Attend the 2016 FSBI annual symposium and present

an oral or poster presentation• Submit to the committee chair a travel report and

photographs for print publication by August 14, 2016• Submit travel receipts to the committee chair by

August 14, 2016 for expense reimbursement• Attend the AFS meeting and IFS committee meeting

in Kansas City, Missouri, USA, August 21-25, 2016• Act as a site host at the Kansas City AFS meeting for

the FSBI Fellow• Serve as the deputy committee chair of the IFS Fellow

Award in 2017 and the committee chair in 2018


FEATURE

Sampling Design for Early Detection of Aquatic Invasive Species in Great Lakes Ports


Joel C. Hoffman U.S. Environmental Protection Agency, Office of Research and Development, Mid-Continent Ecology Division, 6201 Cong-don Blvd., Duluth, MN 55804. E-mail: [email protected]

Joshua SchloesserU.S. Fish and Wildlife Service, Ashland Fish and Wildlife Conservation Office, Ashland, WI

Anett S. Trebitz, Greg S. Peterson, and Michelle GutschU.S. Environmental Protection Agency, Office of Research and Development, Mid-Continent Ecology Division, Duluth, MN

Henry Quinlan U.S. Fish and Wildlife Service, Ashland Fish and Wildlife Conservation Office, Ashland, WI

John R. KellyU.S. Environmental Protection Agency, Office of Research and Development, Mid-Continent Ecology Division, Duluth, MN


We evaluated a pilot aquatic invasive species (AIS) early detection monitoring program in Lake Superior that was de-signed to detect newly introduced fishes. We established survey protocols for three major ports (Duluth-Superior, Sault Ste. Marie, Thunder Bay) and designed an adaptive cycle for routine evaluation of survey performance. Among the three ports, we found both similarities (species richness) and differences (introduced species detectability, species detection efficiency) with respect to AIS survey performance. Despite those differences, our analysis indicated potential for increas-ing detection efficiency at all three ports by exploiting differences in fish assemblages and sampling gears to increase rare species encounters. Using this information in the adaptive cycle, we demonstrate the ability to improve AIS detection efficiency. Our pilot monitoring program with its adaptive cycle of assessment, refinement, and implementation provides a performance-based approach to increase AIS detection efficiency over the course of a survey and within practical re-source constraints.

Diseño de muestreo para la detección temprana de especies acuáticas invasivas en los puertos de los Grandes LagosSe evaluó un programa piloto de monitoreo para la detección temprana de especies acuáticas invasivas (EAI) en el lago Superior, mismo que fue diseñado para detectar peces recién introducidos. Se establecieron protocolos de muestreo en tres puertos importantes (Duluth-Superior, Sault Ste. Marie y Thunder Bay) y se diseñó un circuito adaptativo para evaluar de forma rutinaria el desempeño del muestreo. Con respecto al desempeño del EAI, se encontraron similitudes (riqueza de especies) y diferencias (capacidad de detección de especies introducidas y eficiencia en la detección de especies) entre los tres puertos. Pese a dichas diferencias, los análisis indicaron que existe potencial en los tres puertos para incrementar la eficiencia en la detección de especies si se aprovechan las diferencias entre ensambles ícticos y entre artes de pesca para el muestreo con el fin de incrementar el encuentro de especies raras. Al utilizar esta información en el circuito adap-tativo, se corrobora la habilidad para mejorar la detección del EAI. Nuestro programa de monitoreo piloto, el circuito ad-aptativo de evaluación, su refinamiento e implementación representan un enfoque basado en el desempeño, cuyo objetivo es incrementar la eficiencia en la detección de EAI en el curso de un muestreo y con limitaciones realistas de recursos.

Modèle d'Échantillonnage pour la Détection Précoce des Espéces Aquatique Invasives auxPorts des Grands LacsNous avons évalué un programme pilote de surveillance de la détection précoce d’une espèce aquatique invasive (EAI) [Aquatic invasive species (AIS)] dans le Lac Supérieur, conçu pour détecter les poissons nouvellement introduits. Nous avons établis des protocoles de surveillance pour trois ports majeurs (Duluth-Superieur, Sault Ste. Marie, Thunder Bay), et créé un cycle adaptatif pour l’évaluation habituelle des résultats de la surveillance. Parmi les trois ports, nous avons trouvé des similitudes (diversité des espèces) mais aussi des différences (détéctabilité des espèces introduites, le rendement de la détection des espèces) concernant les résultats de la surveillance des EAI. Malgré ces différences, notre analyse a indiqué le potentiel pour augmenter le rendement de détection dans les trois ports en exploitant les différences dans les communautés de poissons et les équipements d'échantillonnage afin d’augmenter les rencontres des espèces rares. En utilisant cette information dans le cycle adaptatif, nous démontrons la capacité d’améliorer le rendement de détection des EAI. Notre programme pilote avec son cycle adaptatif d’évaluation, amélioration et implémentation fournit une approche basée sur la performance afin d’augmenter le rendement de la détection des EAI sur la durée d’une surveillance et dans les limites des ressources pratiques.

INTRODUCTION

Early detection and rapid response (EDRR) programs are implemented to prevent the establishment and spread of invasive species (Hewitt et al. 2009). Under the recently reauthorized Great Lakes Water Quality Agreement of 2012, a U.S.–Canada aquatic invasive species (AIS) early detection and rapid response program must be established for the Laurentian Great Lakes by 2015 (Inset A). Early detection is important both to

increase the success of rapid response actions (e.g., quarantines, public awareness campaigns, eradication efforts) and to evaluate the success of AIS introduction prevention policies (Simberloff 2003; Lodge et al. 2006; Vander Zanden et al. 2010). Beyond supporting rapid response, early detection AIS monitoring can establish a biodiversity baseline, document species distributions,

and provide information to improve future detection efforts (Lee et al. 2008; Bishop and Hutchings 2011). An ideal early detection program should detect a newly introduced species while it is still rare and geographically restricted (Myers et al. 2000), before its abundance and range has increased to where control is effort intensive, costly, and possibly infeasible (Hulme 2006). Because the probability of detecting an organism is inversely related to its abundance, early detection must be more sensitive to rare species than typical biological surveys designed to monitor plant or animal abundance and distribution (Lodge et al. 2006; Dejean et al. 2012).

To accomplish these aims, monitoring should incorporate regular, long-term surveys that are quantitative and designed using a statistical model (Campbell et al. 2007); EDRR programs, however, rarely implement a monitoring program with all these elements (Hulme 2006). Existing EDRR programs encompass a broad range of jurisdictions, survey designs (active versus passive surveillance), and methods (Table 1). Commonly, EDRR programs are designed to detect watch-list species and are therefore focused on specific habitats near likely introduction points and spread boundaries or, if taxonomically broader, take the form of a database that accepts sighting information but is uninformative concerning survey design or effort (Table 1). Few surveys have been designed to detect a broad taxonomic range of potential invasive species over multiple locations, habitats,

Recognizing that a substantial search effort is needed to find a rare species and that monitoring resources are generally constrained, improving efficiency for early detection is paramount.


Table 1. Examples to represent the range of invasive species EDRR programs in North America. Early detection refers to active surveys specifically aimed at early detection; active detection refers to ongoing surveys, generally for abundance information; passive detection refers to incidental detections.

Program name Target Jurisdiction Partners Detection methods Reference

Eradication program

Asian longhorned beetle Anoplophora glabripennis

Federal

Animal and Plant Health Inspection Service and the Canadian Food Inspection Agency

Early detection: visual survey to locate and destroy infested trees

USFS (2010)

Plan for the prevention, detection, assessment, and management of Asian carps in Michigan waters

Asian carp State University of Notre Dame, An-glers Monitoring Network

Early detection: eDNA

Active: existing fisheries surveys

Passive: public education, state invasive species report-ing system

Clapp et al. (2012)

Integrated pest manage-ment plan for the Surface Water Management Agency of Clackamas County

Vertebrates, inverte-brates, and vegetation

CountyClackamas County depart-ments, districts, and the City of Happy Valley

Passive: County agencies par-ticipate in state and Soil and Water Conservation District reporting system

Guillozet (2012)

Finger Lakes Region Aquat-ic Invertebrate Assessment and Invasive Species Pre-vention Project

Aquatic inva-sive species

Nongov-ernmental organiza-tion

Hobart, William Smith Col-leges, Cayuga County

Passive: public education, stewards placed at boat launches, no formal public reporting system

Cleckner and Go-ranowski (2013)

Invasive Alien Species Partnership Program—Early Detection and Preven-tion Network for Aquatic Invasive Species in the St. Lawrence River

Any new aquatic inva-sive species

Province

Department of Natural Resources and Wildlife of Que-bec, commercial fishers (150 participants)

Active: existing fishery effort

Environ-ment Canada (2010)

INSET A

Great Lakes Water Quality Agreement – 2012, Annex 6

The Parties shall develop and implement programs and other measures to eliminate new introductions of AIS [aquatic invasive species] through a binational prevention-based approach, informed by risk assessments. This approach takes into account that new species may pose a risk to the Great Lakes, even in the absence of scientific certainty.

The Parties, subject to their respective laws and regulations, and in cooperation and consultation with State and Provincial Gov-ernments, Tribal Governments, First Nations, Métis, Municipal Governments, watershed management agencies, other local public agencies, and the Public, shall…within two years of entry into force of this Agreement, develop and implement an early detection and rapid response initiative that:(a) develops species watch lists;(b) identifies priority locations for surveillance;(c) develops monitoring protocols for surveillance;(d) establishes protocols for sharing information;(e) identifies new AIS; and(f) coordinates effective and timely domestic and, when necessary, binational response actions to prevent the establishment of newly detected AIS.

- See more at http://www.ijc.org/en_/GLWQA_Annexes


and collection methods; we are only aware of such programs in Australia currently (Whittle et al. 2013).

Recognizing that a substantial search effort is needed to find a rare species (e.g., at the early stage of introduction; Hoffman et al. 2011) and that monitoring resources are generally constrained, improving efficiency (i.e., number of species detected for a given sampling effort) for early detection is paramount. Species detection (i.e., likelihood of encountering a species) can be increased through additional sampling effort, whether in a short-term intensive phase or spread over multiple years. In contrast, efficiency can be increased by devising an approach to overrepresent rare species in a sample relative to their abundance in the environment (e.g., by targeting specific, possibly difficult to sample habitats; Trebitz et al. 2009) or by amplifying an individual’s signal (e.g., sampling for environmental DNA; Darling and Mahon 2011).

The Great Lakes serve as an excellent case study to evaluate AIS early detection survey design, because they are vulnerable to a wide range of AIS that can spread rapidly once established (Johnson and Padilla 1996; Kornis et al. 2012; Snyder et al. 2014; Pagnucco et al. 2015). We evaluated a pilot AIS early detection monitoring program that was designed to detect newly introduced fishes to Lake Superior and that encompassed three historically and currently high-risk ports for species introductions. Our goal was to assess AIS surveys within each port using a variety of performance metrics and to ask whether we could improve technical capability (rare species detectability) over time at individual locations yet also harmonize survey designs among ports with different environmental qualities and AIS introduction risk. We first compared fish species richness metrics, species detection probabilities, sampling completeness (i.e., observed compared to the estimated species richness), and sampling efficiency among ports. In this context, we asked whether survey performance changes when viewed as separate ports or as a combined data set (to represent a monitoring “network” of ports). We then randomly resampled our survey data to ask whether a similar gear allocation (i.e., percentage of sites sampled by each gear

type) would maximize species richness at each port. Finally, at one of the ports, we used an adaptive approach to determine whether gear allocation changes did in fact improve species detection efficiency. From the pilot program, we draw initial recommendations for a Great Lakes–wide program, recognizing that such a program must accrue information and evolve over time toward a set of not-yet-defined expectations.

METHODSStudy Area

The three ports in the pilot program—Duluth-Superior, Sault Ste. Marie, and Thunder Bay—were chosen because they are at particularly high risk of AIS introductions to Lake Superior. Duluth-Superior and Sault Ste. Marie are among the six Great Lakes new AIS introduction hotspots (Grigorovich et al. 2003). Ballast water discharge has been the most prevalent vector for AIS introduction to the Great Lakes (Ricciardi 2006); Duluth-Superior and Thunder Bay receive a large ballast water discharge volume from both Great Lakes and transoceanic commercial vessels, and both host a variety of AIS (Grigorovich et al. 2003). All three ports are population centers, and both Thunder Bay and Duluth-Superior attract recreational boating and angling activity—factors increasing the introduction risk of AIS associated with commercial trade or recreational boats (Ricciardi 2006; Nathan et al. 2015). Sampling boundaries encompassing roughly equal areas (Duluth-Superior: 44.5 km2; Sault Ste. Marie: 56.3 km2; Thunder Bay: 31.7 km2) were established upriver and downriver of port facilities.

Monitoring ApproachSampling at each port was initiated under an adaptive

monitoring program: a cycle of annual surveillance, evaluation, and improvement (Figure 1). For this study, our focus for improvement was efficiency—how to improve early detection capabilities within practical resource constraints. The goal of initial surveillance was to sample until we encountered nearly all native and introduced fishes known to be present and to acquire supporting environmental data (e.g., habitat characterization,

Figure 1. The adaptive monitoring framework used for the pilot Lake Superior aquatic invasive species early detection monitoring program.


Figure 2. Sampling stations (square for electrofishing, circle for fyke net, triangle for trawl) for Duluth-Superior (2008–2012), Thunder Bay (2010–2012), and Sault Ste. Marie (2010–2012). The bold line represents the boundaries of the designated survey area. Stations sampled in Duluth-Superior in 2006–2007 are reported in Hoffman et al. (2011).

water quality) to evaluate ecological patterns that might facilitate more efficient early detection (Trebitz et al. 2009). A performance evaluation was conducted for each port, and a randomization analysis was used to develop recommendations to increase AIS detection efficiency. For Duluth-Superior, those recommendations were implemented and another cycle of surveillance was conducted, after which we evaluated whether the changes in fact improved survey efficiency.

We began AIS surveillance at Duluth-Superior in 2006 (details in Trebitz et al. 2009; Hoffman et al. 2011; Peterson et al. 2011). We used a random stratified survey design to provide an unbiased selection of sites dispersed throughout the harbor (Stevens and Olsen 2004) with two depth strata based on the depth limit of aquatic vegetation establishment (2.1 m; Angradi et al. 2013). We sampled with three gears that accommodate multiple habitats; gears were assigned to a depth suitable for their operation. Fyke nets and electrofishing sampled a mix of bare and vegetated habitat (≤2 m), and the bottom trawl sampled the dredged navigation channels and thalweg (>2 m). Catch data from the initial 2006 surveillance cycle (n = 95; 25 fyke net, 39 electrofishing, and 31 trawl sites) were then analyzed for survey performance and potential efficiencies. Randomization analysis (detailed description below) revealed that allocating 45–85% of the sites to fyke nets, 10–30% to electrofishing, and 5–25% to trawls most consistently achieved high values for total richness, and introduced species richness, and rare taxa richness simultaneously (Trebitz et al. 2009). Based on this and logistical constraints imposed by the gears, we chose to allocate 40% of the sites to fyke nets, 40% to electrofishing, and 20% to bottom trawls (i.e., a 40:40:20 gear allocation). Starting in 2008, sampling of approximately 50 sites per year using this 40:40:20 gear allocation (again using a random stratified survey design) was implemented at Duluth-Superior and continued through 2012 (Figure 2).

In 2010–2012, Thunder Bay and Sault Ste. Marie were added to the pilot program (Figure 2). For the initial surveillance cycle, 45 sites were selected annually at each port using the same random design and gear-depth allocation as in Duluth-Superior. Habitat varied among ports; within the sampling area, 53.7% of Duluth-Superior, 33.7% of Sault Ste. Marie, and 18.9% of Thunder Bay was less than 2.1 m depth. Stations were divided evenly among the three gears (15 each to fyke net, electrofishing, and bottom trawl) to obtain sufficient sample size to test whether optimal gear allocation is location specific. Plotting of species–effort curves for Duluth-Superior data suggested that >100 samples were necessary to detect ≥95% of the estimated species pool (Hoffman et al. 2011). We chose this 95% target because it tends to be near the asymptote of the species–effort curve, implying a relatively high likelihood of encountering a new, rare species, yet not so close to the asymptote that encountering the next species would require a substantial increase in effort. The AIS surveillance program could sample about 50 sites per year at each port (essentially, five days of effort), and we recommended a three-year evaluation cycle (i.e., available resources defined detection over a three-year period as “early detection”), presuming that combining independent years can reasonably be used as a space–time substitution of effort.

Field MethodsCollected fish were identified to species by fisheries

professionals and tallied; if the identification was uncertain, specimens were vouchered and sent to a collections museum

for verification. We sampled in August–September to maximize the probability of capturing young of year with gears suited to capturing both juvenile and adult fishes. At all three ports, sampling equipment and deployment methods were the same except the timing of electrofishing because daytime fishing at Sault Ste. Marie produced few fish (Table 2).

Data AnalysisFirst, we evaluated performance among the three ports.

Then, for Sault Ste. Marie and Thunder Bay, we conducted a randomization analysis similar to that previously conducted on Duluth-Superior catch data to test whether survey efficiency might be improved by changing gear allocation. We were particularly interested in whether the data supported the


expectation that each port would require a different, tailored sampling approach. Second, we evaluated whether implementing the 40:40:20 gear allocation (i.e., postoptimization) compared to the 2006–2007 initial surveillance effort in Duluth-Superior improved survey performance. We based both evaluations on multiple performance metrics: fish species richness metrics, species-specific detection probabilities, and the species–effort relationship.

The three species richness metrics included were the observed species richness (Sobs), introduced species richness (Snon-native), and rare species richness (Srare-5; those native and nonnative species comprising <5% of the total incidence; Trebitz et al. 2009). We calculated species-specific detection probabilities (hereafter, detectability) for all introduced species as the proportion of samples in which it occurred relative to the total number of samples obtained.

The species–effort relationship describes how quickly species are acquired for a given search effort. Generally, there is a strong, positive correlation between species richness and sampling effort; however, the rate of species acquisition progressively slows to an asymptote, as increasingly rare species require increasing effort to find (Gotelli and Colwell 2001). The asymptote is an estimate of the “true” species pool (hereafter, Sasym). We estimated average species–effort curves and their associated 95% confidence intervals and Sasym values using the EstimateS v8.20 software (Colwell 2006). Whereas a survey encounters species in a sequence that depends on which sites are sampled first, the average increase in species with each additional sample taken can be obtained by randomizing site order (see Hoffman et al. 2011 for technical details). We used sample-based abundance data to maintain the observed patchiness in the catch (Gotelli and Colwell 2001), combining the 2010–2012 data for each port to standardize effort but also combining data from all three ports to gain insight regarding how the different ports scale as a possible monitoring “network.” We measured survey efficiency as the effort required to obtain a coverage of 50%, 95%, or 100% of Sasym (Chao and Jost 2012) using the nonparametric method proposed by Chao et al. (2009). In addition, we compared the difference between the number of species observed (Sobs) and Sasym to estimate how many rare (possibly newly introduced) species were not detected, as well as the effort to detect a newly introduced species (i.e., a very rare species, ≥95% of Sasym).

To identify the optimal allocation of some fixed sampling effort among different combination of gear types with respect to the three richness metrics, we used randomization analyses. Randomizations were based on presence–absence data and a

20-sample draw, chosen because the greatest efficiencies in sampling occur in the early portion of the species–effort curve (Hoffman et al. 2011). Gear combinations were limited to even integers, allowing 66 combinations of 20 samples. For each possible combination, there were 10 random draws; the averages of each of the possible combinations were depicted as contours in two-dimensional triangular space (details in Trebitz et al. 2009).

We hypothesized that gear-based efficiencies are derived from exploiting gear-specific (and thereby habitat-specific) biases in the detectability of different species. To better characterize differences in the patterns of species occurrence among gears and ports, assemblage patterns were summarized by nonmetric multidimensional scaling (NMDS) ordination using Bray-Curtis similarity computed from incidence data. Five unusual samples in which a single species was the entire catch were excluded from the analysis because they strongly influenced the resulting ordination (one Sault Ste. Marie, two Thunder Bay, and two Duluth-Superior samples).

RESULTSPilot Program Evaluation

We detected 11 nonnative fishes, all previously known to occur in Lake Superior (Table 3). Only three species (Alewife Alosa pseudoharengus, Rainbow Smelt Osmerus mordax, Threespine Stickleback Gasterosteus aculeatus) were captured at all ports and with widely varying detectability. Substantially more fish were captured at Duluth-Superior than Sault Ste. Marie or Thunder Bay. Observed species richness (Sobs) was similar among the three ports (36–39 species), but there were differences in the number of nonnative and rare species (Srare-5), as well as singletons (species captured once) and doubletons (species captured twice; Table 4).

Observed species coverage was high for all three ports (Table 4). For each port (and combined), on the cumulative three-year basis (2010–2012), the survey was sampling along the asymptote of the species–effort curve, with Sobs close to or within the lower 95% confidence interval (CI) of Sasym such that there was a nearly 50% probability of sampling Sasym (Figure 3). Thus, a sampling effort similar to that exerted during 2010–2012 would likely detect at least 95% of the estimated species present. The S50% was achieved quickly at Duluth-Superior (just four samples postoptimization) but more slowly at Sault Ste. Marie and Thunder Bay (Figure 3), indicating that new species were acquired at a lower rate at the latter two ports, possibly because many fewer fish were captured per sample at Sault Ste. Marie (mean of 65.9 fish per sample) and Thunder Bay (98.7)

Table 2. Field sampling methods for the pilot aquatic invasive species early detection program by sampling gear, including gear specifications (dimensions, mesh sizes), duration of sampling (“Set”), and deployment methods.

Sampling gear Dimensions, mesh Set Deployment

Fyke net

Frame: 0.9 m (h) × 1.2 m (w) Lead: 15 mMesh: 4.7 mm through-out

>12 hr, overnight

*Sampled in <2 m water depth*Paired nets on both ends of a single lead *Set parallel to shoreline*Substrate type varied with net location (sand, clay, boulder rip rap, vegetation)

Electrofishing vessel

Amperage and voltage determined by local conditions

10 min/station Speed: 3 km/h

*Sampled in <2 m water depth*Daytime: Duluth-Superior, Thunder Bay; Nighttime: Sault Ste. Marie*Effort covered all visible habitat (shoreline, open water) and substrate type (sand, clay, boulder rip rap, vegetation) in sample area

Bottom trawl (Marinovich trawl)

Head rope width: 4.9 mBody mesh: 3.8 cmLiner mesh: 3.125 mm

5 min/stationSpeed: 4 km/h

*Sampled in >2 m water depth*Stations along contour*Sampled dredged navigation channel and thalweg


compared to Duluth-Superior (>300; Table 4). Combining the three ports into a single species–effort curve suggested the network efficiency may be better than its constituent parts. The combined S95% was reached in 225 samples (54% of exerted effort), which was substantially more efficient than Sault Ste. Marie and Thunder Bay individually (102% and 75%, respectively) and similar to Duluth-Superior (45%). Further, on average, 90% of Sasym was sampled annually by the network, though only Duluth-Superior independently attained such coverage.

Randomization analyses revealed that despite differences in fish assemblages and available habitat, there was a similar response to gear allocation mixtures among ports (Figure 4). Contours of the three species richness endpoints examined (Sobs, Snon-native, Srare-5) yielded a similar result, so only Sobs results are shown. In general, allocating less effort to bottom trawling while

increasing fyke net and electrofishing effort tended to yield higher species richness. Allocating more than 40% of sampling effort to trawling resulted in decreased species richness at all locations, but allocating more than 80% of effort to either fyke nets or electrofishing also resulted in decreased richness. For all three ports, allocating the three gears in the 40:40:20 ratio (as implemented at Duluth-Superior) would yield a species richness near or at the maximum observed, though a range of alternative allocations would yield a similar result (Figure 4).

Results from the NMDS were consistent with our interpretation of the contour plots, with gears showing consistent relationships among the three ports even though the fish assemblage varied. Among ports, Duluth-Superior sites generally fell in the upper left quadrant, Thunder Bay sites in the upper right quadrant, and Sault Ste. Marie sites in the lower left quadrant. Within a port, there was little overlap among gears, especially electrofishing and fyke nets, indicating that each contributed different species to the surveillance effort (Figure 5). This confirmed that sampling with multiple kinds of equipment is important to detect rare or unique species and thereby maximize species richness by exploiting gear-specific differences in species detection. Bottom trawls ordinated between and overlapped with both electrofishing and fyke nets, reflecting the observation that few unique species were captured by trawl and accounting for its lower contribution to species richness than the other two gears.

EVALUATION OF DULUTH-SUPERIOR

Detection efficiency improved after optimizing gear allocation for Duluth-Superior. Postoptimization, the number of samples required to reach 95% or 100% of Sasym was reduced by 37% and 58%, respectively (Table 4). Detectability increased slightly for nearly all nonnative species (maximum increase of 0.08), though Brook Silverside Labidesthes sicculus and Threespine Stickleback were not detected in one or the other survey (Table 3). Detectability improved by a factor of 1.4 to 3.0 for rare nonnative species (detectability <0.10 in 2006–2007), whereas it changed little for common nonnative species (probability of detection >0.20). That is, optimization generally increased detectability of rare nonnative species with no or only a slight loss of detectability for common nonnative species (presumably, those readily caught by trawl, the gear for which effort was reduced).

Table 3. Detectability (proportion of samples containing that species) of nonnative fishes captured in each of three ports during the initial surveys and the optimized survey in Duluth-Superior. Fish are arranged by increasing detectability in Duluth-Superior (stage 1). Values are shown for fish that were detected; “NA” indicates that the fish was not detected.

Nonnative species

Initial survey Postoptimization

Duluth-Superior

Sault Ste. Marie

Thunder Bay Duluth-Superior

Brook Silverside <0.01 0.01 NA 0.04

Threespine Stickleback 0.01 0.05 0.07 NA

Alewife 0.03 0.01 0.01 0.07

Freshwater Drum 0.03 0.01 NA 0.09

Rainbow Smelt 0.09 0.02 0.48 0.13

Common Carp 0.13 NA 0.13 0.19

Tubenose Goby 0.16 NA 0.27 0.24

White Perch 0.23 NA NA 0.23

Round Goby 0.39 NA 0.07 0.39

Ruffe 0.46 NA 0.33 0.43

Fourspine Stickleback NA NA 0.13 NA

Table 4. Survey data and sampling completeness statistics for initial surveillance and postoptimized gear allocation by port, including the survey effort by the number of samples and individuals; number of total fish species (Sobs), nonnative species (Snon-native), and rare species (Srare-5; species comprising <5% of the incidence); the number of species for which we captured one (singletons) or two (doubleton) individuals; the estimated number of total fish species (Sasym; in the EstimateS software, the “Chao1” estimator, Chao et al. 2009) and associated 95% CI, and the total effort (samples/individuals) required to detect 50% (S50%), 95% (S95%), or 100% of Sest. At Duluth-Superior, Sasym from the postoptimization survey exactly equals Sobs because the data condition of having two doubleton and one singleton species was met (see Chao et al. 2009).

Initial survey PostoptimizationCombined

Duluth-Superior Sault Ste. Marie Thunder Bay Duluth-Superior

Period 2006 2010–2012 2010–2012 2010–2012 2010–2012

Samples/individuals 95/29,590 135/8,892 135/13,320 150/45,407 420/67,619

Sobs/Snon-native/Srare-5 35/9/6 36/5/17 37/8/12 39/9/9 57/11/28

Singletons/doubletons 2/1 4/4 2/3 1/2 2/2

Sasym (95% CIs) 36 (35–43) 37 (36–46) 37 (37–42) 39 (39–39) 57 (57–63)

Effort:Sasym 352/109,265 337/22,196 219/21,627 150/45,407 862/138,796

Effort:S95% 105/31,897 138/9,116 101/9,965 67/20,282 225/36,222

Effort:S50% 5/1,557 13/856 7/691 4/1,211 11/1,771


Figure 3. Species-effort curves for the three ports (A-C; 2010-2012) and all three ports combined (D; 2010-2012 only), including the num-ber of observed species (Sobs), as well as the average species-effort curve (solid line) and its 95% confidence interval (CI; dashed lines). The various levels of effort required to sample the estimated species total (Sasym; error bars show 95% CI), as well as 50% (S50%) and 95% (S95%) of Sasym are indicated. A site is a location where a sampling gear was deployed once.

Figure 4. Contour plot of total species richness accumulated over 20 station-sets as a function of the mix of sampling gears. The axes and upper right diagonal bound the possible station–gear combi-nations, so percentage fyke netting stations equals 100 minus the percentage of electrofishing (y-axis value) and bottom trawl (x-axis value) stations. As an aid to integrating information across the figure panels, the small square highlights the region associated with 40% fyke netting, 40% electrofishing, and 20% trawling effort (i.e., the 40:40:20 allocation).


DISCUSSION

We found both similarities (observed species richness, relative species coverage) and differences (introduced species detectability, species acquisition efficiency) with respect to AIS survey performance among the three ports. Notably, our perception of survey performance changed with respect to scale in that combining ports into a network-level analysis revealed detection efficiencies not observed at the port level. Despite survey performance differences and differences with respect to introduction vectors, aquatic habitat, and fish assemblages among the ports, the randomization analysis indicated that changes to gear allocation are expected to improve survey efficiency and that a single gear allocation strategy might simultaneously maximize species richness at all three ports. Using this information in the adaptive cycle of evaluation, recommendation, and implementation demonstrated an ability to improve AIS detection efficiency through time. Here, we discuss whether we found evidence to treat ports differently with respect to design and performance metrics, design-based limits to data interpretation, and lessons learned from the pilot program to inform Great Lakes–wide AIS early detection monitoring.

For all three ports surveyed, a similar level of initial effort (~50 stations per year, a ca. one- to two-week effort for a management agency) provided a useful starting point for surveillance. At each port, about 30 fish species (~80% of Sasym) were detected in 50 samples, suggesting that allocating effort equally among ports (of a roughly equal area) provides a reasonable starting basis for a lake-wide program. Initially, annual surveillance was likely missing six to eight species; at least 100 samples (2–3 years of effort) were needed to detect >95% of species present (Sasym).

With three years of data to optimize surveillance design (i.e., Duluth-Superior results), we substantially improved detection efficiency. The survey detected nearly 95% of Sasym annually and >99% of Sasym on a two-year basis, which equates to an effort reduction of about 50% to detect most (at least 95%) of the fish assemblage (Sasym). The species–effort curve and NMDS analyses provide some insight into how these efficiencies were achieved. By biasing the design to emphasize some types of sampling (i.e., fyke nets and electrofishing), we increased effort in certain types of places (e.g., relatively complex, structured habitat) where initial sampling yielded the greatest number of species and most distinct catches, thereby increasing efficiency. Yet, we cannot eliminate any gear entirely because as the NMDS illustrated, each gear contributed unique species within a port.

The combined species–effort curve revealed that the network of ports acquired more species and was more efficient (detected species faster) than the constituent port surveys. This appears to be a landscape-scale effect somewhat analogous to the

The goal of an adaptive program is to allow better management and program functional-ity against the backdrop of ecological change and technological and resource constraints, rather than to provide incrementally better estimates of performance.

Figure 5. Two-dimensional NMDS ordination (stress = 0.17) based on incidence data from 2010 to 2012 for all three ports, with sampling gear indicated (T: bottom trawl; E: electrofishing; N: fyke netting). The three ports were ordinated together but are displayed as three panels for clarity.


within-port optimization; efficiency was improved because the individual port surveys each contributed unique species but also different detection probabilities for shared species (i.e., a species that is rare at one port is common at another port, so detection efficiency is improved by combining the two). This suggests that including habitats of differing environmental qualities at both the within- and among-port scales provides useful variability in species presence and detectability that can be exploited for increased efficiency. Of course, we did not test the effect of changing the effort allocation among ports at the network scale. Reducing the total effort at any specific location can potentially delay detection of a new AIS, giving it more time to expand and potentially increasing control and management costs (Epanchin-Neill et al. 2012), though this risk can be managed using the adaptive monitoring cycle. Performance metrics provide information regarding the risk of AIS nondetection for a given level of effort at each location surveyed; managers can respond by establishing achievable goals with respect to AIS early detection and then adapting the monitoring design to meet these goals.

The randomization analysis showed remarkably strong convergence in the optimized gear ratio among ports despite differences in introduction vectors, area sampled, fish assemblage, and habitat distribution. The analysis thus demonstrated that gear-specific biases can be exploited. There may be environmental variables other than gear (i.e., depth) such as temperature strata (cool versus warm water fishes) that can be used to improve efficiency; this is readily testable within the adaptive monitoring cycle. As conceived, with each new cycle, managers ask a design question with respect to species detectability and detection efficiency and then identify the required supporting data for performance evaluation.

Though we found the performance metrics chosen to provide useful information for survey evaluation, there remain limits to design-based knowledge that should be recognized. For example, the surveillance used three sampling gears that target juvenile and adult fish and was conducted in late summer. Other sampling targets (e.g., fish larvae, invertebrates), methods (e.g., gill nets, seines, plankton nets, benthic samplers), and timing (e.g., spreading effort over multiple seasons) may provide new avenues for improving efficiency. We anticipate that using this adaptive cycle in other Great Lakes ports could yield different strategies to optimize AIS surveillance. In addition, detectability estimates are constrained by the effort exerted (e.g., it is not possible to determine whether a fish occurs at a 1:100,000 detection probability until 100,000 fish are sampled), so the concept of what constitutes “rare,” as well as which species are rare, will evolve over time in a given port. Ultimately, the goal of an adaptive program is to allow better management and program functionality against the backdrop of ecological change (i.e., new vectors or AIS) and technological and resource constraints (Ringold et al. 1996), rather than to provide incrementally better estimates of performance. Thus, potential adaptations ought to emphasize consideration of survey designs (Campbell et al. 2007) or technologies that might markedly improve AIS early detection (e.g., DNA-based detection; Darling and Mahon 2011).

Any AIS early detection surveillance program must contend with the trade-off between optimizing for broad taxonomic coverage versus species-specific detectability (Bishop and Hutchings 2011). We designed the monitoring program to be nonspecific (i.e., to detect any nonnative fish), rather than to tar-

get one or few species, enabling us to use rare species detection to inform survey power (i.e., species–effort relationship) but not to directly estimate survey power. In contrast, when targeting a single species, it is possible to maximize its detection probability (e.g., by using a tailored sampling method or targeting a specific habitat) and estimate the likelihood of a false negative detection (Whittle et al. 2013) but at the risk of reduced detectability of nontarget species. Some mix of strategies may be feasible in the context of an AIS early detection program. At stake is whether the survey has sufficient power to detect a newly introduced and by definition rare AIS. There is no straightforward method to estimate the probability that a newly introduced species remains undetected. The advantage of the species–effort curve is that it provides context for what effort is sufficient to catch rarely en-countered species regardless of their identity (Chao et al. 2009; Chao and Jost 2012) and allows comparisons among locations and even ecosystems (Gotelli and Colwell 2001). In addition, species–effort metrics allow the monitoring agency to report a lack of new detection with an associated level of confidence, thereby helping to avoid the perception that new AIS were not detected simply for lack of effort.

Although AIS early detection programs can benefit from existing fisheries index surveys, the goals and requirements of an AIS early detection survey are quite distinct. Efficient AIS early detection surveys should be designed to increase the opportunity to encounter rare species, maximize differences in species composition among sampling events (i.e., among gears and locations), and minimize repetitive sampling that does not yield new taxa. These aims run counter to the repeatability and standardization that fisheries index surveys typically seek. Further, correct identification to species is important to recognize new AIS (Bishop and Hutchings 2011) as well as rare species because they drive the performance evaluation by influencing the shape of the species–effort curve. Again, this is in contrast to the typical fisheries index survey, which tends to treat rare species as “noise” and minimize time spent identifying species that contribute little to trend indices.

Our adaptive monitoring pilot provided a performance-driven approach to investigate, implement, and evaluate potential efficiencies for AIS early detection (within design-based limits to our knowledge). As such, we perceive it is broadly applicable to the problem of invasive species early detection monitoring. Treating the various ports using a generalized approach appeared to be adequate, and we found little evidence of needing to tailor sampling effort, analyses, and approaches to individual ports. However, we have only investigated one of many areas where efficiencies may be gained. A lake- or basin-wide program would include locations with a broader array of environments and introduction risk than found within a port and, as such, tailoring survey sampling effort and equipment to locations may improve survey performance. As the Great Lakes AIS early detection monitoring program develops and performance criteria are established, our strategy can provide a foundation for adaptive monitoring, acknowledging that timely reporting to managers will be critical to AIS detection risk management and that new designs and technologies must be evaluated and incorporated along the way.

ACKNOWLEDGMENTS

Field sampling was conducted by the U.S. Fish and Wildlife Service and U.S. Environmental Protection Agency in partnership with the Ontario Ministry of Natural Resources,


the Fond Du Lac Tribe Resource Management Agency, and the 1854 Treaty Authority, with supporting individuals including N. Bogyo, B. Borkholder, C. Butterworth, S. Chong, T. Corry, G. Czypinski, A. Edwards, S. Greenwood, T. Kaspar, R. McNeely, D. Montgomery, K. Schmidt, B. Sederberg, I. Senecal, and J. Van Alstine. We thank E. O’Malia for GIS assistance, A. Olsen for the survey design, M. Starry for map production, and. B. Nagle (University of Minnesota Bell Museum) for outside expertise on fish taxonomy. The views expressed in this article are those of the authors and do not necessarily reflect the views or policies of the U.S. Fish and Wildlife Service or the U.S. Environmental Protection Agency.

REFERENCESAngradi, T. R., M. S. Pearson, D. W. Bolgrien, B. J. Bellinger, M. A.

Starry, and C. Reschke. 2013. Predicting submerged aquatic veg-etation cover and occurrence in a Lake Superior estuary. Journal of Great Lakes Research 39:536–546.

Bishop, M. J., and P. A. Hutchings. 2011. How useful are port surveys focused on target pest identification for exotic species manage-ment? Marine Pollution Bulletin 62:36–42.

Campbell, M. L., B. Gould, and C. L. Hewitt. 2007. Survey evalua-tions to assess marine bioinvasions. Marine Pollution Bulletin 55:360–378.

Chao, A., R. K. Colwell, C.-W. Lin, and N. J. Gotelli. 2009. Sufficient sampling for asymptotic minimum species richness estimators. Ecology 90:1125–1133.

Chao, A., and L. Jost. 2012. Coverage-based rarefaction and extrapo-lation: standardizing samples by completeness rather than size. Ecology 93:2533–2547.

Clapp, D. F., J. L. Mistak, K. M. Smith, and M. A. Tonello. 2012. Pro-posed 2010 plan for the prevention, detection, assessment, and management, of Asian carps in Michigan waters. Michigan De-partment of Natural Resources, Fisheries Division, Special Re-port 60, Lansing.

Cleckner, L., and K. Goranowski. 2013. Finger Lakes Institute Wa-tercraft Steward Program. Available: flisteward.wordpress.com/about. (January 2014).

Colwell, R. K. 2006. EstimateS software. Available: viceroy.eeb.uconn.edu/EstimateS. (May 2014).

Darling, J. A., and A. R. Mahon. 2011. From molecules to manage-ment: adopting DNA-based methods for monitoring biologi-cal invasions in aquatic environments. Environmental Research 111:978–988.

Dejean, T., A. Valentini, C. Miquel, P. Taberlet, E. Bellemain, and C. Miaud. 2012. Improved detection of an alien invasive species through environmental DNA barcoding: the example of the American bullfrog Lithobates catesbeianus. Journal of Applied Ecology 49:953–959.

Environment Canada. 2010. Invasive Alien Species Partner-ship Program. Available: www.ec.gc.ca/nature/default.asp?lang=En&n=B008265C-1. (January 2014).

Epanchin-Neill, R. S., R. G. Haight, L. Berec, J. M. Kean, and A. M. Liebhold. 2012. Optimal surveillance and eradication of invasive species in heterogeneous landscapes. Ecology Letters 15:803–812.

Gotelli, N. J., and R. K. Colwell. 2001. Quantifying biodiversity: pro-cedures and pitfalls in the measurement and comparison of spe-cies richness. Ecology Letters 4:379–391.

Grigorovich, I. A., R. I. Colautti, E. L. Mills, K. Holeck, A. G. Ballert, and H. J. MacIsaac. 2003. Ballast-mediated animal introductions in the Laurentian Great Lakes: retrospective and prospective analyses. Canadian Journal of Fisheries and Aquatic Sciences 60:740–756.

Guillozet, P. 2012. Integrated pest management plan for the Surface Water Management Agency of Clackamas County, Clackamas County Service District No. 1, and the City of Happy Valley. Avail-able: www.clackamas.us/wes/documents/IntegPestManPlan.pdf. (January 2014).

Hewitt, C. L., R. A. Everett, and N. Parker. 2009. Examples of current international, regional and national regulatory frameworks for

preventing and managing marine bioinvasions. Pages 335–352 in G. Rilov and J. A. Crooks, editors. Biological invasions in ma-rine ecosystems. Springer-Verlag, Berlin.

Hoffman, J. C., J. R. Kelly, A. S. Trebitz, G. S. Peterson, and C. W. West. 2011. Effort and potential efficiencies for aquatic non-na-tive species early detection. Canadian Journal of Fisheries and Aquatic Sciences 68:2064–2079.

Hulme, P. E. 2006. Beyond control: wider implications for the man-agement of biological invasions. Journal of Applied Ecology 43:835–847.

Johnson, L. E., and D. K. Padilla. 1996. Geographic spread of exotic species: ecological lessons and opportunities from the invasion of the zebra mussel Dreissena polymorpha. Biological Conser-vation 78:23–33.

Kornis, M. S., N. Mercado-Silva, and M. J. Vander Zanden. 2012. Twen-ty years of invasion: a review of Round Goby Neogobius mela-nostomus biology, spread and ecological implications. Journal of Fish Biology 80:235–285.

Lee, H., D. A. Reusser, J. D. Olden, S. S. Smith, J. Graham, V. Burkett, J. S. Dukes, R. J. Piorkowski, and J. McPhedran. 2008. Integrat-ed monitoring and information systems for managing aquatic invasive species in a changing climate. Conservation Biology 22:575–584.

Lodge, D. M., S. Williams, H. J. MacIsaac, K. R. Hayes, B. Leung, S. Reichard, R. N. Mack, P. B. Moyle, M. Smith, D. A. Andow, J. T. Carlton, and S. McMichael. 2006. Biological invasions: recom-mendations for U.S. policy and management. Ecological Appli-cations 16:2035–2054.

Myers, J. H., D. Simberloff, A. M. Kuris, and J. R. Carey. 2000. Eradica-tion revisited: dealing with exotic species. Trends in Ecology and Evolution 15:316–320.

Nathan, L. R., C. L. Jerde, and M. L. Bundy. 2015. The use of environ-mental DNA in invasive species surveillance of the Great Lakes commercial bait trade. Conservation Biology 29:430–439.

Pagnucco, K. S., G. A. Maynard, S. A. Fera, N. D. Yan, T. F. Nalepa, and A. Ricciardi. 2015. The future of species invasions in the Great Lakes–St. Lawrence River basin. Journal of Great Lakes Research 41(Suppl. 1):96–107.

Peterson, G. S., J. C. Hoffman, A. S. Trebitz, C. W. West, and J. R. Kelly. 2011. Establishment patterns of non-native fishes: lessons from the Duluth-Superior harbor and lower St. Louis River, an invasion-prone Great Lakes coastal ecosystem. Journal of Great Lakes Research 37:349–358.

Ricciardi, A. 2006. Patterns of invasion in the Laurentian Great Lakes in relation to changes in vector activity. Diversity and Distribu-tions 12:425–433.

Ringold, P. L., J. Alegria, R. L. Czaplewski, B. S. Mulder, T. Tolle, and K. Burnett. 1996. Adaptive monitoring for ecosystem management. Ecological Applications 6:745–747.

Simberloff, D. 2003. How much information on population biology is needed to manage introduced species? Conservation Biology 17:83–92.

Snyder, R. J., L. E. Burlakova, A. Y. Karatayev, and D. B. MacNeill. 2014. Updated invasion risk assessment for Ponto-Caspian fishes to the Great Lakes. Journal of Great Lakes Research 40:360–369.

Stevens, D. L., and A. R. Olsen. 2004. Spatially-balanced sampling of natural resources. Journal of the American Statistical Associa-tion 99:262–278.

Trebitz, A. S., J. R. Kelly, J. C. Hoffman, G. S. Peterson, and C. W. West. 2009. Exploiting habitat and gear patterns for efficient detection of rare and non-native benthos and fish in Great Lakes coastal ecosystems. Aquatic Invasions 4:651–667.

USFS (U.S. Forest Service). 2010. Forest disturbance processes—Asian longhorned beetle. Available: www.nrs.fs.fed.us/distur-bance/invasive_species/alb. (January 2014).

Vander Zanden, M. J., G. J. A. Hansen, S. N. Higgins, and M. S. Kornis. 2010. A pound of prevention, plus a pound of cure: early detec-tion and eradication of invasive species in the Laurentian Great Lakes. Journal of Great Lakes Research 36:199–205.

Whittle, P. J. L., R. Stoklosa, S. Barrett, F. C. Jarrad, J. D. Majer, P. A. J. Martin, and K. Mengersen. 2013. A method for designing com-plex biosecurity surveillance systems: detection non-indigenous species of invertebrates on Barrow Island. Diversity and Distri-butions 19:629–639.


Frank J. RahelDepartment of Zoology & Physiology, Program in Ecology, University of Wyoming, 1000 E. University Ave., Laramie, WY 82071. E-mail: [email protected]

Changing Philosophies of Fisheries Management as Illustrated by the History of Fishing Regulations in Wyoming

FEATURE


Inland fisheries management began in the United States in the 1800s with a focus on fish as food and the use of stocking to create new fisheries and replenish depleted stocks. In the early 20th century, recreational fishing came to the forefront and regulations limiting the number and size of fish that could be harvested were enacted. Major trends in the regulation of recreational fisheries included a reduction in creel limits, more complexity in the application of regulations, increas-ingly restrictive use of baitfish, and limitations on competitive fishing tournaments. In the latter part of the 20th century, fisheries managers embraced a broader perspective that included conservation of native species and control of invasive species. These changes in regulations reflect the evolution of fisheries management philosophy along pathways emphasiz-ing fishing for sustenance, fishing for recreation, and, most recently, biodiversity management. This evolution is illustrated by the history of angling regulations in Wyoming.

Cambio en la filosofía del manejo de pesquerías ilustrado por la historia de las regulaciones pesqueras en WyomingLas pesquerías continentales en los EE.UU. iniciaron en el siglo XIX y consideraban a los peces como fuente de alimento y, por medio de la estabulación, servían para crear nuevas pesquerías y para recuperar poblaciones agotadas. A inicios del siglo XX, apareció la pesca recreativa y entraron en vigor las regulaciones que limitaban el número y tamaño de los peces capturados. Las principales tendencias en cuanto a la regulación de las pesquerías recreativas incluyeron límites a las cantidades de especies capturadas, mayor complejidad en la aplicación de regulaciones, uso cada vez más restrictivo de carnadas y mayores limitaciones en los torneos de pesca. Hacia finales del siglo XX, los administradores pesqueros adoptaron una visión más integral que incluyó la conservación de especies nativas y el control de especies invasoras. Estos cambios en las regulaciones se reflejaron en la evolución de la filosofía del manejo, haciendo énfasis en la pesca de subsistencia, pesca recreativa y, más recientemente, en el manejo de la biodiversidad. Esta evolución es ilustrada por la historia de las regulaciones a la pesca con caña en Wyoming, EE.UU.

L’évolution des Philosophies de Gestion de la Pêche comme l’Illustre l’Histoire desRéglementations de pêche dans le WyomingLa gestion de la pêche dans les eaux intérieures a commencé aux États-Unis dans les années 1800, avec un accent sur le poisson en tant que nourriture et l’utilisation des réserves afin de créer de nouvelles ressources halieutiques et renouveler les stocks diminués. Au début du 20ème siècle, la pêche de loisir s’est hissée au premier plan et des réglementations ont été adoptées limitant le nombre et la taille du poisson qui pouvait être récolté. Les tendances majeures de la réglementa-tion concernant la pêche de loisir incluaient la réduction des limites, plus de complexité dans l'application des réglementa-tions, de plus en plus de restrictions quant à l’utilisation du poisson-appât, et des limitations concernant les compétitions de pêche sportive. Vers la fin du 20ème siècle, les responsables de la pêche ont adopté une vision plus large comprenant la conservation des espèces indigènes et le contrôle des espèces invasives. Ces modifications des réglementations re-flètent l'évolution des philosophies de gestion de la pêche en suivant les courants qui ont mis l’accent sur la pêche comme nourriture, la pêche comme loisir et, plus récemment, la gestion de la biodiversité. Cette évolution est illustrée par l’histoire des réglementations de la pêche à la ligne dans le Wyoming, États-Unis d’Amérique.

INTRODUCTION

Regulations are a major aspect of inland fisheries management in the United States. Purchasers of recreational fishing licenses often receive a detailed booklet outlining when and where angling is allowed, which species can be pursued, and how many and what size fish can be harvested. The types and complexity of fishing regulations have changed greatly since the first restrictions on fish harvest in the early 1900s. These changes in regulations reflect the evolution in management philosophy along three main lines: fishing for sustenance, fishing for recreation, and management of biodiversity (i.e., conservation of native species and control of invasive species; Figure 1). Although the time course and relative importance of regulatory changes may differ among states, they represent a common set of responses to challenges facing managers of inland recreational fisheries.

I use the history of fishing regulations in Wyoming to illustrate the evolution of fisheries management philosophies in the United States. Regulations in Wyoming from 1869 through 1938 were obtained from a database of Wyoming statutes (HeinOnline 2014) and from Wiley (1993). From 1939 through 2015, fishing regulations were published in brochures available from the Wyoming Game and Fish Department (WGFD; Cheyenne, Wyoming).

SUSTENANCE FISHING

Fisheries management in the United States began in the late 1800s with a focus on the use of fish for sustenance. Federal

and state fisheries agencies were established with the aim of introducing species to create new fisheries or replenishing exploited wild stocks with hatchery fish. On June 10, 1872, the U.S. Congress passed a bill authorizing the U.S. Fish Commission, the forerunner of the U.S. Fish and Wildlife Service, to commence fish culture and stocking (Moffitt et al. 2010). Stocking by private individuals also became common. For example, by the 1870s, members of the California Ornithological and Piscatorial Acclimatizing Society were already at work introducing eastern Brook Trout Salvelinus

Figure 1. Fisheries management in the United States has evolved along pathways emphasizing sustenance fishing, recreational fishing, and biodiversity management.


fontinalis into California waters (Halverson 2010). An 1879 article in the Laramie newspaper noted the efforts of anglers who caught trout in Colorado and, in violation of Colorado game laws, surreptitiously transported them into Wyoming for release into the Laramie River, which had no native trout species. Even noted ichthyologist David Starr Jordan recommended stocking nonnative catfishes into tributaries of the lower Colorado River because “the whole great basin of the Colorado contains, excepting the trout, no fish of even second-rate character as food for man” (D. S. Jordan 1889:6).

Because of unregulated harvest and habitat destruction, many fish populations were considered to be in deplorable condition by the late 19th century. Barkwell (1883:6) noted that many Wyoming streams were nearly exhausted of a once bountiful supply of food fish and included among the causes “barbarous methods of taking fish such as the use of giant powder and poisonous drugs.” Market hunting—that is, harvesting large numbers of animals to be sold for profit—further contributed to the decimation of wildlife and fish populations (Blair 1987). Regulations were soon enacted to stop the wasteful overexploitation of fish while still allowing for sustenance harvest. In Wyoming, the first law pertaining to fishing was passed by the Territorial Legislature in 1869 and stated that hook and line was the only legal means by which fish could be harvested, thus outlawing the use of dynamite and poisons that allowed large numbers of fish to be harvested at once (Glafcke 1876:363). In 1875, a second law stated that only wildlife and fish in the amount “necessary for human subsistence governed in amount and quantity by the reasonable necessities of the person” could be harvested (Glafcke 1876:362). This law was difficult to enforce and market hunting was more directly targeted by an 1899 law that made the sale of game fish illegal in Wyoming (Van Orsdel and Chatterton 1899). During the late 1800s, other laws mandated fishways at dams, required

screening of irrigation ditches, and prohibited sawdust or mining waste from being discharged into waterways. Similar laws were passed throughout North America during this period (Moffitt et al. 2010).

RECREATIONAL FISHINGHarvest and Size Limits

It became increasingly apparent by the late 1800s that stocking alone could not compensate for the continuing decline in fish populations. Although legislation had outlawed the use of dynamite, nets, and poisons to catch fish, overharvest from hook-and-line fishing remained a problem in Wyoming and elsewhere (Figure 2). Michigan’s first superintendent of fisheries described a similar situation due to the lack of harvest regulations (Jerome 1875, cited in Clark et al. 1981):

That waters once abounding with fish can become barren by excessive, or ill-timed, or barbarous fishing, or all together, is too obviously, painfully true. … Laws, too, prescribing closure times and regulating the utensils and methods of capture, whether by seine or weir, or spear or hook, grow out of the very necessities of the case. … It is absence or nonobservance of these laws that has depleted many a stream and river, pond and lake, of all their finny wealth and beauty.

To stem the depletion of fish populations, states began enacting regulations in the early 20th century to limit the harvest of game fish. In Wyoming, the first daily creel limit was established in 1899 at 20 pounds of game fish (generally understood to refer to trout). This remained in effect until 1931 when a limit of 15 pounds or 30 game fish was established (Table 1). The limit was further reduced to 15 pounds or 20 game fish in 1937. In Michigan, the first creel

Figure 2. Trout caught by fly fisherman at West Thumb, Yellowstone Lake, Yellowstone National Park, Wyoming, 1897 (Nolan 1983).


limit for trout was 50 per day in 1903 (Clark et al. 1981). In Minnesota, the earliest daily creel limits were 25 for Walleye Sander vitreus, Northern Pike Esox Lucius, and Largemouth Bass Micropterus salmoides in 1910 (Cook et al. 2001). Even in national parks where wildlife preservation is a major goal, creel limits were high; for example, 20 Yellowstone Cutthroat Trout Oncorhynchus clarkii bouvieri per day from Yellowstone Lake in 1908 (Gresswell and Varley 1988). By today’s standards, these creel limits seem incredibly high, but one can only imagine the resentment they must have engendered among anglers accustomed to harvesting fish

without limit. Daily creel limits were continually reduced during the 20th century, and today anglers can keep few fish, especially among large-bodied game species (Table 1).

Regulations have become increasingly complicated as managers attempted to maximize the fishing experience based on biological constraints and angler preferences. The first fishing regulations in Wyoming specified limits for “game fish,” but the taxa that comprised game fish were not defined until 1945 (Hunt et al. 1945). The number of taxon-specific harvest regulations has increased greatly over the past century (Table 2). A major reason for the increased complexity of regulations

Table 1. Changes in game fish daily creel limits in states for which historical summaries of angling regulations are available. Creel limits are expressed as number of fish unless collective fish weight in pounds is specified. Data from the 2010s are from management agency websites. Historical data are from following sources: Wyoming (current study), Great Smoky National Park (Kulp and Moore 2005), Montana (Montana Fish, Wildlife and Parks, Helena, Montana), North Dakota (North Dakota Game and Fish Department, Bismarck, North Dakota), Michigan (Clark et al. 1981; Diana and Smith 2008), Minnesota (Cook et al. 2001), Yellowstone National Park (Gresswell and Varley 1988), Ohio (Carey Knight, Ohio Department of Natural Resources, personal communication), Maine (R. M. Jordan 2001), Pennsylvania (Weber et al. 2010), and Utah (Drew Cushing, Utah Division of Wildlife Resources, personal communication).

Period

Wyoming

Yellowstone Lake,

Yellowstone National

Park

Great Smoky

Mountains National

Park

Maine Michigan Minnesota

Trout Black bass Walleye

Yellowstone Cutthroat

TroutTrout Black

bass Trout Northern Pike Walleye Northern

Pike

Large-mouth Bass

1900s 20 pounds None None 20 Unknown None 50 None None None None

1910s 20 pounds None None 20 Unknown 25 35 25 25 25 25

1920s 20 pounds None None 20 Unknown 25 15 5 15 25 15

1930s 30 None None 10 10 25 15 5 8 10 6

1940s 12 or 20a 20 20 5 10 25 15 5 8 3 6

1950s 12 or 20a None 12 or 20a 5 5 15 10 5 6 3 6

1960s 10 None 10 3 5 12 5 5 6 3 6

1970s 6 or 10a None 6–10a 3 4 8 5 5 6 3 6

1980s 6 10 10 2 5 5 5 5 6 3 6

1990s 6 6 6 2 5 1 or 3a 5 5 6 3 6

2000s 6 6 6 0 5 1 or 3a 5 2 6 3 6

2010s 3 or 6b 6 6 0 5 1 or 3a 5c 2 6 3 6

PeriodMontana North

Dakota Ohio Lake Erie Pennsylvania Utah

Trout Black bass Walleye/Sauger Trout Yellow Perch Trout Trout

1900s None None None 25 None None 10 pounds

1910s None None None 15 None None 10 pounds

1920s 40 40 40 15 None 25 10 pounds

1930s 15 15 15 15 None 10–20a 10 pounds

1940s 15 15 pounds 15 15 None 10 10 pounds

1950s 10 15 15 5 None 8 10

1960s 10 15 15 5 None 8 10

1970s 5 or 10d 10–15a 10–15a 5 None 8 8

1980s 5 or 10d 5–10a 5–10a 5 50 8 8

1990s 5 or 10d 5 5 5 30 8 8

2000s 5 5 5 3 30 5 4

2010s 5 5 5 3 30 5 4

aVaries by watershed.bLimit is three for streams and six for lakes.cLimit is five for streams but ranges from 1 to 5 in lakes.dLimit is five for streams and 10 for lakes.


is a large increase in geographic and temporal exceptions to state or regional regulations (Table 2). For example, in 1931, there was a single regulation on the number of game fish that could be harvested in Wyoming and a statewide fishing season from April 1 to November 30 except for a later start in two counties. By 1939, dozens of water bodies had exceptions to the statewide fishing season or creel limits. In 1948, Wyoming was divided into five major drainage areas, each with its own fishing regulations, although local exceptions to season and creel limits were common. By 1950, angler age restrictions appeared for various waters. By 1980, gear restrictions allowing only artificial flies or lures were put into place on several streams (Table 2). Catch-and-release requirements appeared by 1990. Of particular note is the advent of mandatory kill requirements in 2014 for game fish species deemed undesirable in particular waters. This regulation is discussed further in the section below on invasive species.

Also contributing to the increased complexity of angling regulations was the trend to adjust creel limits, size restrictions, and gear restrictions depending upon local productivity, fishing pressure, and the public’s desire for particular fishing

experiences. Consider the North Platte River in central Wyoming. In 1947, the statewide creel limit of 12 trout applied to the entire river except from the city of Casper to the inlet of Seminoe Reservoir where the creel limit was five trout. In 1988, size limits and gear restrictions differed among three sections of the river, although the creel limit was consistent at six fish. By 2014, the river was divided into 10 sections with different creel limits, size limits, and gear restrictions. Regulations were least restrictive in reservoirs where hatchery fish supported a harvest-oriented fishery. By contrast, sections of the river that supported fisheries renowned for large wild trout were managed with low creel limits and a flies-and-lures-only restriction to facilitate the catch-and-release fishery practiced by many anglers using those areas.

Baitfish RegulationsSeining had been outlawed in 1869 in Wyoming to protect

game fish, but that law did not allow for collection of baitfish. As a concession to anglers who wanted to use live fish as bait, the law was changed in 1931 to allow seining for baitfish provided that seining was not done in waters frequented by game fish. The general trend since then has been increasing restrictions on

Table 2. The increasing complexity of angling regulations in Wyoming. Gamefish categories refers to the number of game fish taxa that have their own creel limits. An asterisk denotes the listing of waters with an exception to the state or area-wide regulations for fishing season, creel or size limit, age of anglers, gear restrictions, catch-and-release fishing only, or mandatory kill of undesirable species.

Year Gamefish categories

Fishing season

Creel or

size limit

Angler age

restriction

Gear restriction

Catch and

release

Mandatory kill Comments

1900 1 — — — — — — Gamefish not defined but assumed to be trout species.



1930 1 * — — — — — Gamefish not defined but assumed to be trout species.

1940 1 * * — — — — Gamefish not defined except in a few waters with limits for certain trout species.

1950 2 * * * — — — Regulations for general game fish and Burbot Lota lota.

1960 5 * * * — — —

Regulations for general game fish, Mountain Whitefish Prosopium williamsoni, Brook Trout Salvelinus fontinalis, Burbot, and Grayling Thymal-lus arcticus.

1970 5 * * * — — — Same as 1960.

1980 8 * * * * — —

Regulations for general game fish, Mountain Whitefish, Brook Trout, Burbot, Grayling, Walleye/Sauger Sander canadensis, black bass Micropterus spp., and Northern Pike.

1990 10 * * * * * —

Regulations for general trout, Brook Trout, Moun-tain Whitefish, black bass, Walleye/Sauger, catfish, Burbot, Northern Pike/Tiger Musky Esox lucius × E. masquinongy, sturgeon Scaphirhynchus plato-rynchus, and panfish.

2000 10 * * * * * — Same as 1990.

2010 11 * * * * * *Regulations for general trout, Brook Trout, Moun-tain Whitefish, black bass, Walleye, Sauger, catfish, Burbot, Northern Pike/Tiger Musky, sturgeon, and panfish.

2015 12 * * * * * *Regulations for general trout, Brook Trout, Lake Trout Salvelinus namaycush , Mountain White-fish, black bass, Walleye, Sauger, catfish, Burbot, Northern Pike/Tiger Musky, sturgeon, and panfish.


where baitfish can be used, which species can be used as bait, and how baitfish can be procured (Table 3). A major reason for baitfish regulation was to prevent the introduction of non-game species that might be detrimental to game fish (Remmick 1982). In 1946, WGFD noted that anglers had introduced suckers (Catostomidae) into numerous Wyoming lakes, where they grew too big to provide forage and were thought to eat the eggs of game fish (Spratt 1946). It is interesting that, in many cases, the game fish of concern were not themselves native species but had been stocked in mountain lakes or low-elevation reservoirs that lacked fish species of interest to anglers.

To reduce the chances that baitfish would become established outside their native drainages, WGFD in 1951 required that baitfish had to be used in the waters where they had been collected. The use of live baitfish was banned in the Green River and Bear River drainages in 1971 to protect highly valued trout fisheries from the negative effects of illegally introduced non-game species. Also in that year, anglers possessing live baitfish had to produce a sales receipt or copy of their seining permit to verify the origin of the fish.

A major change occurred in 1974 when the use of live baitfish was restricted to a subset of drainages east of the Continental Divide in Wyoming (Table 3). Fisheries management in those drainages had expanded from a focus on trout to include coolwater and warmwater nonnative species such as Walleye and black bass Micropterus spp. that had become naturalized in these systems. Using live baitfish was a preferred method of angling for these species. To regulate the sale of baitfish more effectively, vendors were required to obtain a bait dealer’s license, allow inspection of their facilities, and maintain records of sales. However, there were no restrictions on the sources or species of baitfish that could be imported from other states.

Baitfish regulations were further tightened in 1996 when only Fathead Minnows Pimephales promelas and Golden Shiners Notemigonus crysoleucas could be imported by baitfish

dealers (Table 3). Dealers were required to notify WGFD 72 hours prior to importation so that baitfish shipments could be inspected. Anglers could keep commercially purchased live baitfish for 10 days, after which the fish had to be killed. Concerns that baitfish shipments from out-of-state sources might be contaminated with other fish species led to a ban on the importation of all baitfish in 1999. Since then, there have been further restrictions on how baitfish can be obtained, and in 2012 vendors could purchase only Fathead Minnows from licensed hatcheries for sale as live baitfish (Table 3).

Fishing TournamentsIn Wyoming, competitive fishing tournaments have been

around since 1983 when the Saratoga Chamber of Commerce sponsored an ice fishing derby to stimulate business during the slow winter season. Early tournaments focused on trout, but competitive fishing did not become widespread until 1990 with the advent of events focused on Walleye. In response to the growing number of tournaments, WGFD published the first regulations for fishing contests in 1990. A fishing contest was defined as “any competitive angling event conducted on waters in the State of Wyoming for the purpose of awarding prizes, or for personal gain or promotional consideration.” Such events required written approval by WGFD at least 10 days prior to the event.

The trend has been for regulation of fishing tournaments to become more prescriptive. In 1992, a fishing contest was more precisely defined as “any event for catching game fish from waters open to public use where an entry or participation fee of $5.00 or more is charged per angler, 50 or more anglers participate on a given date, or total prizes exceeding $1,000 in cash or merchandise are awarded.” Contests had to be approved 30 days prior to the event, and sponsors were required to submit a summary report. In addition, written approval was required to release fish in a live-release fishing contest. This was done to ensure that live-release contests would only be held under

Table 3. History of regulations regarding the use of baitfish in Wyoming.

Year Regulations1869–1930 No regulations regarding use of baitfish.

1931 Legal to seine for baitfish except in waters frequented by game fish.

1948 Illegal to have in possession while fishing any “live bait fish or rough fish.” Legal to seine for minnows except in waters frequented by game fish.

1951 Legal to use live baitfish only in waters where they were collected.

1969 Same as 1951 except that a permit was required to seine baitfish.

1971Legal to use live baitfish only in waters where they were collected. Exception: live baitfish not allowed in Green River and Bear River drainages. Permit required to collect bait fish. Persons with live bait in possession must have receipt or permit verifying origin of fish. Ban on importation of all live fish except with authorization.

1974Use of live baitfish restricted to selected drainages. Where legal, live baitfish have to be collected in drainage where they will be used and a seining permit is required. Commercial sale of live baitfish allowed through a baitfish dealer license and any species can be imported from out of state. Anyone using live baitfish must have sales receipt or seining permit.

1996Where legal, live baitfish must be collected in drainage where they will be used and a seining permit is required. Only Fathead Minnow Pimephales promelas and Golden Shiner Notemigonus crysoleucas can be imported from out of state by baitfish dealers. Baitfish shipments subject to inspection. Commercially purchased live baitfish can only be kept for 10 days.

1999 Importation of all baitfish from out of state banned.

2000 Fathead Minnow and Golden Shiner can be used statewide wherever live baitfish allowed.

2004 Possession of live Brook Stickleback Culaea inconstans prohibited. Illegal to import amphibians, reptiles, crustaceans, or mollusks as live bait. Mollusks and crustaceans caught in Wyoming can be used as live bait only in waters where collected.

2008 Commercially purchased live baitfish can be kept for 15 days except that Fathead Minnow and Golden Shiner can be kept for 30 days.

2012 Commercial baitfish dealers can purchase only Fathead Minnows from licensed hatcheries and sell them anywhere live bait is legal.


conditions conducive to survival of fish that had been held in live wells and then weighed prior to being released. Because fishing tournaments in Wyoming are concentrated on relatively few reservoirs, conflicts arose between tournament anglers and members of the public who did not appreciate having to share their favorite fishing spots with numerous competitive fishers. To lessen this conflict, a “Special Fishing Contest Provision” was adopted in 2006 whereby certain water bodies would have at least two weeks that were free from fishing contests each year. The most recent regulation requires anglers to harvest the Walleye they catch in tournaments after July 1 because high water temperatures in summer result in high mortality of Walleye released after being weighed (Hoffman et al. 1996).

BIODIVERSITY MANAGEMENTProtecting Native Species

Throughout the first half of the 20th century, there was little recognition by anglers or fisheries managers as to whether game fish species were native or nonnative. Consequently, nonnative species such as Rainbow Trout O. mykiss were widely stocked throughout the United States to provide fish for the creel (Halverson 2010). In addition, it was common to combine similar taxa for regulatory purposes; hence, creel and size limits were often set for categories such as “trout” that did not distinguish between native and nonnative species. But in the 1970s, interest in conservation of declining native trout species in the western United States began to develop (Behnke and Zarn 1976). In Wyoming, conservation efforts began for several subspecies of native Cutthroat Trout O. clarkii. Early efforts involved habitat improvements, but in 1984, in response to population declines, some Wyoming streams with Cutthroat Trout were closed to fishing. Protection for Cutthroat Trout was expanded in 1990 when numerous streams throughout the state were converted to catch-and-release fishing. Harvest regulations were further tightened in 2008 to allow only two of the six total trout creel limit to be Cutthroat Trout within their native range in Wyoming. Another native game fish that has received regulatory attention in Wyoming is Sauger Sander canadensis. Prior to 2008, the creel limit was six fish for any combination of Sauger and Walleye, the latter being nonnative to Wyoming. The two species were separated in 2008, and the creel limit for Sauger

was set at only two fish, whereas the creel limit for Walleye remained at six fish.

Passage of the Endangered Species Act of 1973 and a growing awareness of the importance of protecting biodiversity in all of its forms led natural resource management agencies to add non-game fishes and aquatic taxa such as mussels and crayfish to their management responsibilities (Schramm and Hubert 1999). An important first step in management of non-game species is to recognize their value in ecosystems. Early fishing regulations in Wyoming defined creel and season limits for game fish and listed qualifying species by their common name. All other species were classified as “rough, coarse, or non-game species.” Starting in 1973, species not specifically listed as game fish were simply referred to as “non-game fish.” An effort to remove terms such as “rough fish” or “trash fish” from the lexicon of fisheries biologists was also occurring elsewhere in the United States (Martin 1976; Woodling 1985). This change in terminology for fish not utilized as game species was an important event in the evolution of fisheries management philosophy because it indicated a turning point in how non-game species were viewed by biologists. Use of terms such as rough or coarse fish by a state management agency reinforces the public’s perception that these species have little value and therefore makes it difficult to engender support for conservation of non-game taxa that are declining such as species of suckers or minnows (Cyprinidae).

Regulations can promote fish conservation goals by creating protected areas where no fishing or bait collecting is allowed. For example, Kendall Warm Springs in northern Wyoming was closed to fishing and bait collecting starting in 1978 to protect the endemic Kendall Warm Springs Dace Rhinichthys osculus thermalis.

Managing Invasive SpeciesInvasive species are nonnative species whose introduction

to an ecosystem is likely to cause environmental or economic harm or harm to human health (Kolar et al. 2010). In Wyoming, concern about invasive fish species was evident by the middle of the 20th century when managers noted the harmful efforts of illegal introductions on sport fisheries (Spratt 1946). Later, concern about invasive species was extended to include their harmful effects on native, non-game species, such as Bluehead

Figure 3. Evolution of angling regulations in relation to the control of aquatic invasive species in Wyoming.


Sucker Catostomus discobolus, Flannelmouth Sucker C. latipinnis, and Roundtail Chub Gila robusta (discussed in Bezzerides and Bestgen 2002). Fishing regulations can play a role in preventing introduction of invasive species and in controlling their population size after they have become established. A regulation passed in 1937 made it illegal to stock fish in Wyoming without a permit (Figure 3), but release of live baitfish persisted. This led to increasingly restrictive regulations on the use of baitfish beginning in 1951 (Table 3). Although baitfish regulations were largely enacted to prevent harm to game fish, they had the added benefit of protecting native non-game fishes from invasive species. In an effort to prevent fish stocking by the public, a regulation was enacted in 1976 that made it illegal to transport live fish, even if the fish had been legally harvested. In 2004, possession of Brook Stickleback Culaea inconstans was outlawed and it became illegal to import amphibians, reptiles, crustaceans, or mollusks as live bait.

The year 2006 marked a new development in the use of regulations to manage invasive fish species in Wyoming. For the first time, game fish species that had been illegally introduced to waters where their presence was deemed detrimental were targeted for removal by anglers. In the Green River and Bear River drainages of southwestern Wyoming, the creel limit for Walleye and Burbot Lota lota, considered invasive species in these drainages, was liberalized to 25 fish and all individuals captured had to be killed immediately (Figure 3). This meant that anglers fishing for other species or who captured small fish were required to kill Walleye and Burbot even if they did not intend to consume them. Unfortunately, this regulation conflicted with a 1988 regulation that made it illegal to “take and leave, abandon or allow any game fish … to intentionally or needlessly go to waste.” The mandatory kill aspect of this regulation was removed in 2008, and the creel limit was made unlimited. In the meantime, biologists worked to have the Wyoming Legislature pass a law that allowed game fish to be reclassified as non-game fish in waters where they were considered to be invasive. As a result, in 2014, Burbot, Yellow Perch Perca flavescens, Northern Pike, and Walleye were classified as non-game species in the Green River, Bear River, and Little Snake River drainages of western Wyoming with unlimited creel limits and a mandatory kill designation. In that same year, Walleye in Buffalo Bill Reservoir outside of Yellowstone National Park was classified as a non-game species with an unlimited creel limit and mandatory kill designation.

Three other regulations were enacted recently in an effort to reduce the introduction of nonnative species in Wyoming waters. In 2004, it became illegal to import amphibians, reptiles, crustaceans, or mollusks as live bait. In addition, mollusks and crustaceans collected in Wyoming could only be used as bait in the waters where they were collected. In 2010, the fine for illegal fish stocking was increased from $1,000 to $10,000 to provide a stronger deterrent to anglers releasing live baitfish or surreptitiously stocking game fish into a water body. In 2012, it became mandatory for all boats to stop at inspection stations operated by WGFD along major highways entering Wyoming. The main objective was to prevent boaters from bringing nonnative mussels Dreissena spp. and aquatic plants into Wyoming. Because live-wells in boats are inspected, the program also prevents live fish from being transported into Wyoming.

DISCUSSION

Fishing regulations in Wyoming have evolved in response to three major philosophies guiding inland fishing management (Figure 1). The regulatory trends seen in Wyoming occurred in other parts of the United States as well (Clark et al. 1981; Cook et al. 2001; Isermann and Paukert 2010). The earliest regulations were minimal and reflected a generous allowance for harvesting fish as food. Methods such as dynamite or poisons that killed large numbers of fish without regard to species or size were outlawed. These were analogous to the practice of market hunters who slaughtered big game animals in large and unsustainable numbers in the 19th century. But as fishing developed into a recreational pastime in the late 1800s and early 1900s in the United States, there was increasing concern about the quality of the fishing experience and less tolerance for anglers who justified large fish catches on the basis of sustenance. Although subsistence fishing is not a major activity in Wyoming, it still occurs in some areas of North America (Moffitt et al. 2010).

After establishing hook and line as the only legal means of fishing, fisheries managers turned their attention to creel and minimum size limits at the beginning of the 20th century. However, the earliest creel limits were excessively high and unsustainable, necessitating a continual trend toward reduced harvest limits (Table 1). Regulations have also been adjusted to reflect variation in biological productivity and angler desires. Fisheries biologists have experimented with a variety of regulatory innovations involving gear restrictions, catch and release, and mandatory kill of undesired species. As a result, fishing regulations now include numerous exceptions to the general regulations (Paukert et al. 2001, 2007; Isermann and Paukert 2010). This complexity and the occasional exclusion of entire classes of anglers from some fisheries has led to some backlash; hence, fisheries agencies now try to balance the need for biological specificity and legal clarity with social equity and simplicity (Thurow and Schill 1994; Cooke et al. 2013).

Regulations regarding the use of live baitfish have become more restrictive in Wyoming (Table 3) and throughout North America. Currently, nine Canadian provinces and seven U.S. states either ban or greatly restrict the use of live baitfish (Drake and Mandrak 2014). Earlier concerns dealt with the effects of baitfish on sportfish, especially species such as Common Carp Cyprinus carpio and suckers that obtain large sizes and compete with game species (Remmick 1982). Today, there is increasing concern about nonnative baitfish as disease vectors and as invasive species. In surveys of bait dealers’ tanks, nontarget species are often present, including species known to be invasive (Drake and Mandrak 2014). Despite public education programs and regulations prohibiting release of live bait, the practice remains common. For example, 30% of anglers illegally released live baitfish in Ontario and 65% of anglers did so in Maryland (Kilian et al. 2012; Drake and Mandrak 2014). Because a segment of the fishing public appears refractory to efforts to stop the release of live bait, increasingly stringent regulations on use of baitfish appear likely in the future.

In the latter part of the 20th century, fisheries managers added preservation of biodiversity to their ongoing efforts to enhance recreational fishing. Often this meant stringent harvest limitations on game fish of conservation concern such as Cutthroat Trout and Bull Trout S. confluentus (Erhardt and Scarnecchia 2014). Although closing a fishery may be the most effective way to recover declining fish species, it disenfranchises


anglers whose expenditures and enthusiasm for outdoor recreation contribute to conservation programs (Cooke et al. 2014). Therefore, catch and release or reduced creel limits are a compromise that protects species while maintaining the support of recreational anglers.

Increasingly, management agencies are also restricting collection of non-game species of conservation concern. In Colorado, it is illegal to take 24 non-game fish species, including species considered undesirable or rough fish in the past. This trend will undoubtedly continue as agencies recognize more species as being of conservation concern.

One of the more interesting shifts in regulations involves mandatory kill of invasive fishes that are highly valued in other parts of their distribution. A good example is the Burbot in Wyoming. This species is native and of conservation concern in the Missouri River drainage of Wyoming. Creel limits are restricted and efforts are underway to increase Burbot populations in this area. By contrast, Burbot is not native across the Continental Divide in the Colorado River drainage. There, Burbot, along with Walleye, Northern Pike, and Yellow Perch, are considered invasive species and must be killed when caught by anglers. This duality of species being considered desirable in some areas but undesirable in other areas will increase as fisheries managers use all avenues to stem the tide of illegal fish introductions. Unlimited harvest limits and mandatory kill regulations have also been used to help control invasive populations of Lake Trout S. namaycush in Yellowstone Lake and other Western U.S. waters (Martinez et al. 2009). Even where mandatory kill regulations do not have a major impact on the abundance of invasive species, they send an important message to the public that illegal stocking is harmful and will not be rewarded by managing the invasive species as a desirable game fish (Johnson et al. 2009).

SPECULATIONS ON THE FUTURE OF FISHING REGULATIONS IN THE UNITED STATES

How will management philosophies and fishing regulations be affected by human demographic trends or changing beliefs held by the public? Will a growing human population put more recreational angling pressure on fish populations, or will angling pressure decline because of “nature deficit disorder”; that is, reduced participation in nature-based recreation in an increasingly urbanized society (Pergams and Zardiac 2008; Arlinghaus et al. 2015). A declining number of anglers coupled with increased voluntary catch-and-release practices would mitigate the need for the increasingly restrictive harvest limits that characterized the 20th century (Table 1). Conversely, some forms of angling, such as fly-fishing, have grown in popularity and increased the pressure on some fisheries. Expanded catch-and-release regulations would seem to be the best way to ensure a quality experience in these fisheries. Segmentation of fisheries into areas with different angling regulations will also likely increase as managers try to satisfy the expectations of different angler groups (Thurow and Schill 1994).

A regulation that may become more widespread is a closure on angling for trout during periods of low stream flows and warm water temperatures. High temperatures and accompanying low oxygen concentrations are stressful to coldwater fish and increase the likelihood of mortality due to angling. To prevent mortality, streams in Montana and Yellowstone National Park are closed to fishing between 2 p.m. and midnight when maximum water temperatures reach at least 22.8°C for three

consecutive days. Fisheries biologists in Colorado can close waters to fishing when average daily water temperatures exceed 22.2°C or daily minimum oxygen levels are below 5 ppm. The WGFD recommends that anglers stop fishing for trout when water temperatures exceed 21.1°C, but this has not been codified into a regulation. With many studies predicting increased stream water temperatures due to global warming, regulations mandating fishing closures due to high water temperatures will likely become more common.

Changing attitudes about animal welfare may challenge current thinking about fishing regulations. Although many anglers consider catch and release to be the epitome of ethical fishing, some people believe that the only justification for subjecting fish to the stress and pain of being caught with a hook is to provide human sustenance (Arlinghaus et al. 2007). Strong opposition to catch-and-release fishing exists in Germany and Switzerland, where fish of legal size that are captured must be harvested (Arlinghaus et al. 2007). In the future, anglers and managers might find it difficult to justify catch-and-release fishing in the face of opposition from the animal welfare movement (Arlinghaus et al. 2012).

Regulations related to invasive species will likely increase. Johnson et al. (2009) suggested that the low fines associated with illegal fish stocking should be increased given the high cost of controlling invasive species. This happened in Wyoming when fines for illegal stocking were increased 10-fold in 2010. Johnson et al. (2009) also suggested that a reward system be implemented to encourage people to turn in individuals who illegally stock fish. This would be an interesting throwback to the earliest days of fisheries management in Wyoming when the Territorial Legislature established a $50 fine for violation of game laws, with one-quarter of the fine awarded to the informer (Glafcke 1876). Regulations regarding which species are illegal to possess will almost certainly become more prescriptive to make it easier to prosecute violators. Fishing regulations in Wyoming in 2012 stated that it was illegal to stock or possess aquatic invasive species, but there was no list of species considered to be invasive. In 2014, this lack of clarity was eliminated when 14 taxa (including six species of fish) were listed as aquatic invasive species. Such lists will likely be expanded as agencies identify more species they want to prevent from becoming established within their jurisdictions.

The primary focus of inland fisheries management changed from providing sustenance to providing recreational opportunities in the early 20th century. But could the pendulum swing back to put more emphasis on fish as a food resource? Changing immigration patterns in the United States mean that more people come from cultural backgrounds that emphasize consumptive uses of fish, including taxa such as carps and suckers that were not historically targeted by recreational anglers (Arlinghaus et al. 2007). In addition, might economic hardship and a trend to utilize local food sources (the locavore movement; Tidball et al. 2013) contribute to increased interest in harvesting fish for consumption? Angling regulations have tracked changing philosophies of fisheries management over the past 150 years, and it will be interesting to see how regulations will evolve in response to future challenges to provide recreational and sustenance fishing opportunities while protecting biodiversity in aquatic ecosystems.

ACKNOWLEDGMENTS

A. L. Conder, J. Muhollem, M. A. Smith, D. D. Miller, K. R. Gelwicks, D. J. Zafft, and M. D. Stone provided information on


the history of angling regulations in Wyoming. B. Geddings and D. Skaar provided historical information on fishing regulations in Montana. W. A. Hubert, D. D. Miller, D. J. Schill, A. W. Walters, R. W. Wiley, and an anonymous reviewer provided insightful comments on the article.

REFERENCESArlinghaus, R., S. J. Cooke, J. Lyman, D. Policansky, A. Schwab, C.

Suski, S. G. Sutton, and E. B. Thorstad. 2007. Understanding the complexity of catch-and-release in recreational fishing: an in-tegrative synthesis of global knowledge from historical, ethical, social, and biological perspectives. Reviews in Fisheries Science 15:75–167.

Arlinghaus, R., A. Schwab, C. Riepe, and T. Teel. 2012. A primer on anti-angling philosophy and its relevance for recreational fisher-ies in urbanized societies. Fisheries 37(4):153–164.

Arlinghaus, R., R. Tillner, and M. Bork. 2015. Explaining participation rates in recreational fishing across industrialized countries. Fish-eries Management and Ecology 22:45–55.

Barkwell, M. C. 1883. Biennial report of the state fish commissioner of Wyoming for the years 1882 and 1883. Wyoming Game and Fish Department, Cheyenne.

Behnke, R. J., and M. Zarn. 1976. Biology and management of threat-ened and endangered western trouts. USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, General Technical Report RM-28, Fort Collins, Colorado.

Bezzerides, N., and K. R. Bestgen. 2002. Status review of Roundtail Chub Gila robusta, Flannelmouth Sucker Catostomus latipinnis, and Bluehead Sucker Catostomus discobolus in the Colorado River basin. Colorado State University, Department of Fishery and Wildlife Biology, Larval Fish Laboratory Contribution 118, Fort Collins.

Blair, N. 1987. The history of wildlife management in Wyoming. Wyo-ming Game and Fish Department, Cheyenne.

Clark, R. D., Jr., G. R. Alexander, and H. Gowing. 1981. A history and evaluation of regulations for Brook Trout and Brown Trout in Michigan streams. North American Journal of Fisheries Manage-ment 1(1):1–14.

Cook, M. F., T. J. Goeman, P. J. Radomski, J. A. Younk, and P. C. Jacobson. 2001. Creel limits in Minnesota: a proposal for change. Fisheries 26(5):19–26.

Cooke, S. J., Z. S. Hogan, P. A. Butcher, M. J. W. Stokesbury, R. Ra-ghavan, A. J. Gallagher, N. Hammerschlag, and A. J. Danylchuk. 2014. Angling for endangered fish: conservation problem or conservation action? Fish and Fisheries doi: 10.1111/faf.12076

Cooke, S. J., C. D. Suski, R. Arlinghaus, and A. J. Danylchuk. 2013. Voluntary institutions and behaviours as alternatives to formal regulations in recreational fisheries management. Fish and Fish-eries 14:439–457.

Diana, J. S., and K. Smith. 2008. Combining ecology, human de-mands, and philosophy into the management of Northern Pike in Michigan. Hydrobiologia 601:125–135.

Drake, D. A. R., and N. E. Mandrak. 2014. Ecological risk of live bait fisheries: a new angle on selective fishing. Fisheries 39(5):201–211.

Erhardt, J. M., and D. L. Scarnecchia. 2014. Population changes after 14 years of harvest closure on a migratory population of Bull Trout in Idaho. North American Journal of Fisheries Manage-ment 34:482–492.

Glafcke, H. 1876. The compiled laws of Wyoming. Available: heinon-line.org/HOL/Index?collection=sstatutes. (March 2014).

Gresswell, R. E., and J. D. Varley. 1988. Effects of a century of human influence on the Cutthroat Trout of Yellowstone Lake. Pages 45–52 in R. E. Gresswell, editor. Status and management of interior stocks of Cutthroat Trout. American Fisheries Society, Sympo-sium 4, Bethesda, Maryland.

Halverson, A. 2010. An entirely synthetic fish. How Rainbow Trout beguiled America and overran the world. Yale University Press, New Haven, Connecticut.

HeinOnline. 2014. State statutes: a historical archive. Available: heinonline.org/HOL/Index?collection=sstatutes. (March 2014).

Hoffman, G. C., D. W. Coble, R. V. Frie, F. A. Copes, R. M. Bruch, and K. K. Kamke. 1996. Walleye and Sauger mortality associated with live-release tournaments on the Lake Winnebago system, Wisconsin. North American Journal of Fisheries Management 16:364–370.

Hunt, L. C., W. Jack, and L. J. O’Marr. 1945. Wyoming compiled stat-utes. The Bobbs-Merrill Company, Indianapolis, Indiana. Avail-

able: heinonline.org/HOL/Index?collection=sstatutes. (March 2014).

Isermann D. A., and C. A. Paukert. 2010. Regulating harvest. Pages 185–212 in W. A. Hubert and M. C. Quist, editors. Inland fisheries management in North America, 3rd edition. American Fisheries Society, Bethesda, Maryland.

Johnson, B. M., R. Arlinghaus, and P. J. Martinez. 2009. Introduced species—are we doing all we can to stem the tide of illegal fish stocking? Fisheries 34(8):389–394.

Jordan, D. S. 1889. Report of explorations in Colorado and Utah dur-ing the summer of 1998, with an account of the fishes found in each of the river basins examined. Bulletin of the United States Fish Commission 9:1–40.

Jordan, R. M. 2001. Black bass management plan. State of Maine Department of Inland Fisheries and Wildlife, Augusta.

Kilian, J. S., R. J. Klauda, S. Widman, M. Kashiwagi, R. Bourquin, S. Weglein, and J. Schuster. 2012. An assessment of a bait industry and angler behavior as a vector of invasive species. Biological Invasions 14:1469–1481.

Kolar, C. S., W. R. Courtenay, Jr., and L. G. Nico. 2010. Managing un-desired and invading fishes. Pages 213–259 in W. A. Hubert and M. C. Quist, editors. Inland fisheries management in North Amer-ica, 3rd edition. American Fisheries Society, Bethesda, Maryland.

Kulp, M. A., and S. E. Moore. 2005. A case history of fishing regu-lations in Great Smoky Mountains National Park: 1934–2004. North American Journal of Fisheries Management 25:510–524.

Martin, R. G. 1976. Philosophy of sport fisheries management. Fisher-ies 1(6):8–10, 29–30.

Martinez, P. J., P. E. Bigelow, M. A. Deleray, W. A. Fredenberg, B. S. Hansen, N. J. Horner, S. K. Lehr, R. W. Schneidervin, S. A. Tolen-tino, and A. E. Viola. 2009. Western Lake Trout woes. Fisheries 34(9):424–442.

Moffitt, C. M., G. Whelan, and R. Jackson. 2010. Historical perspec-tives on inland fisheries management in North America. Pages 1–41 in W. A. Hubert and M. C. Quist, editors. Inland fisheries management in North America, 3rd edition. American Fisheries Society, Bethesda, Maryland.

Nolan, E. E. 1983. Northern Pacific views: the railroad photography of F. Jay Haynes, 1876–1905. Montana Historical Society Press, Helena.

Paukert, C. P., J. A. Klammer, R. B. Pierce, and T. D. Simonson. 2001. An overview of Pike regulations in North America. Fisheries 26(6):6–13.

Paukert, C. P., M. McInerny, and R. Schultz. 2007. Historical trends in creel limits, length-based limits, and seasonal restrictions for black basses in the United States and Canada. Fisheries 32(2):62–72.

Pergams, O. R. W., and P. A. Zaradic. 2008. Evidence for a fundamen-tal and pervasive shift away from nature-based recreation. Pro-ceedings of the National Academy of Sciences 105:2295–2300.

Remmick, R. 1982. Live bait can kill a fishery. Wyoming Wildlife 46(1):30–31.

Schramm, H. L., Jr., and W. A. Hubert. 1999. Ecosystem management. Pages 111–123 in C. C. Kohler and W. A. Hubert, editors. Inland fisheries management in North America, 2nd edition. American Fisheries Society, Bethesda, Maryland.

Spratt, R. W. 1946. State of Wyoming, biennial report of the Wyo-ming Game and Fish Commission, 1945–1946. Wyoming Game and Fish Department, Cheyenne.

Thurow, R. F., and D. J. Schill. 1994. Conflicts in allocation of wild trout resources: an Idaho case history. Pages 132–140 in R. Barn-hart, B. Shake, and R. H. Hamre, editors. Wild trout V: wild trout in the 21st century. Trout Unlimited, Arlington, Virginia.

Tidball, K. G., M. M. Tidball, and P. Curtis. 2013. Extending the loca-vore movement to wild fish and game: questions and implica-tions. Natural Sciences Education 42:185–189.

Van Orsdel, J. A., and F. Chatterton. 1899. Revised statues of Wyo-ming. Available: heinonline.org/HOL/Index?collection=sstatutes. (March 2014).

Weber, R., R. T. Green, J. Arway, R. S. Carney, and L. Young. 2010. History of the management of trout fisheries in Pennsylvania. Pennsylvania Fish and Boat Commission, Division of Fisheries Management, Bellefonte.

Wiley, R. W. 1993. Wyoming fish management, 1869–1993. Wyoming Game and Fish Department, Fish Division, Administrative Re-port, Cheyenne.

Woodling, J. 1985. Colorado’s little fish: a guide to the minnows and other lesser known fishes in the state of Colorado. Colorado Di-vision of Wildlife, Denver.

BOOK REVIEW


Anadromous salmon in the Pacific and Atlantic oceans are iconic species that have been the subjects of numerous natural science texts that narratively describe their common plight. Among my favorites is Bruce Brown’s Mountain in the Clouds: A Search for Wild Salmon. Reading this book is one reason that I became passionate about wild salmon conservation. However, there has been a gap in this narrative for the eastern United States, where Atlantic Salmon Salmo salar

is only part of a mosaic of ocean-going species. Running Silver is an intriguing read that fills this void. The book weaves the story of the plight of Northwest Atlantic diadromous fish with source material as varied as Thoreau and Atlantic States Marine Fisheries Commission management actions. Despite the diverse sources, this narrative achieves the author’s goal of explaining the demise of this complex of species while trying to restore biological and cultural awareness of what has been lost … and could be restored. John Waldman’s passion and dedication to the sea run fish community (both the fish and the people) is apparent in each of the book’s 20 relatively short 10-20 page chapters. Although they are woven together into a compelling story, most of the chapters are largely independent, allowing easy reading in small doses. The text transitions between number heavy text needed to explain harsh facts of lost abundance and hectares of concrete that block Atlantic Ocean rivers to very personal narrative observations of fish and river systems. I particularly appreciated the frequent conversational text where Waldman related discussions and observations of river keepers, managers, and scientists. The reader feels as if they are at the table or the river’s edge.

The first four chapters set the stage and describe the fish species and river systems of the Atlantic realm. Chapter 3, The Seasonal Parade, was a favorite. Waldman uses the annual sequence of migration to introduce the reader to both the primary diadromous species and some that may be better characterized as euryhaline wanderers. This chapter was concise yet nuanced in its description of diversity within and among diadromous fish in their ecology and habitat use. And who knew there were anadromous whitefish!

Running Silver: Restoring Atlantic Rivers and their Great Fish Migrations

BOOK REVIEW

John Waldman. Lyons Press. Guilford, CT. 2013. 284 pages. US$26.95 (hardcover), $12.50 (Kindle).

Reviewed by: John F. Kocik NOAA-Fisheries Northeast Fisheries Science Center, 17 Godfrey Drive - Suite 1, Orono, ME 04473. E-mail: [email protected]

It is at this point that the author inserts the first of two interludes. It is 1600, and a female American Shad Alosa sapidissima is beginning her trip to the ocean in a free-flowing river system. Waldman’s insights into the sights, sounds, and hazards faced by this single fish in massive schools, follow her journey to sea and back. This paints a stark contrast with Interlude II (after Chapter 18) that follows a fish’s journey in 2013. Together these two interludes effectively provide a “fish-eye view” of the modern challenges of river and ocean life.

Between the interludes, Chapters 5 through 18 thoroughly and informatively cover the primary challenges to sea run fish, what are often called the four H’s in salmon circles—Habitat, Hydropower, Harvest, and Hatcheries. I enjoyed the way these hazards were covered with stories, case studies, and narratives by passionate professionals and fish advocates. We learn more about river herring, sturgeon, Striped Bass Morone saxatilis, etc., as we explore the challenges the fish face and the people that work to save them. The story is balanced and scientifically accurate; we see and feel the impacts of development on these fish populations. However, it is not a preachy and scolding narrative. The book is an honest and compelling true story that needed to be assembled in one book. In addition to the four H’s, two modern and emerging threats are well documented and described—climate change and ecosocial anomie. With climate change, he describes the realities facing fish at the southern edge of their range and the changes and shifts that face the more broadly distributed species such as river herring. He also introduces a new hurdle: ecosocial anomie—a challenge caused when managers lack accurate baselines and the public has forgotten the utility, and beauty, of these species. This hurdle seems very difficult to address. These six challenges to diadromous fish are mentioned and reinforced throughout the book—this solidifies these challenges and puts the reader on notice that solutions are needed. Beyond the bad news, there are stories of hope such as the Striped Bass story. It is comforting to know that when science finds an answer to population decline and managers are able to prescribe a direct solution to the problem, the fish will respond.

The final two chapters provide a description of optimistic changes in attitudes and river connectivity that have occurred in the last decade and finally a way forward towards better stewardship of these resources. The author’s prescription for recovery is outlined in 10 steps. While the challenges seem insurmountable, a book like this is essential to starting the conversation and starting a fire. As Brown’s 1982 salmon book did for me, I suspect that John Waldman’s book will foster an obsession for river herring and other sea run fish in some readers that might have a career ahead in saving Atlantic rivers and their great fish migrations.


The Hutton Junior Fisheries Biology Program Do you want to Inspire young high school students?

Do you desire to share your Passion for fisheries?

Participate in promoting Diversity & Inclusion?

If you Answered Yes! YES!

Then Apply NOW!

AMERICAN FISHERIES SOCIETY

Contact: Cynthia Oboh, Educational Program Coordinator, via [email protected] or [email protected]

Apply Online NOW! @ hutton.fisheries.org

Mentors hold the key to Engage, Inspire, and Enrich the next generation of fisheries professionals.

To help with this, AFS staff are committed to working with mentors during the Hutton Program.

The Hutton Junior Fisheries Biology Program is a paid summer internship and mentoring program for high school juniors and seniors interested in pursuing the disciplines of fisheries science, marine biol-ogy, and STEM related fields, and is sponsored by the American Fisheries Society (AFS). From the be-ginning of the program in 2001 to 2015, the program has offered scholarships to 572 Hutton Scholars with the help of 270 mentors in 137 host and financially sponsoring institutions.

Selected students known as “Hutton Scholars,” are matched and mentored by a fisheries professional to enjoy an eight-week hands-on fisheries science summer experience in a marine and/or freshwater set-ting. Hutton Scholars receive a $4,000 scholarship award.

Our Mission:

⛵ To increase diversity within the fisheries professions.

Our Vision:

The principle goal of the Hutton Program:

⛵ Is to stimulate interest in careers in fisheries science and management among groups un- derrepresented in the fisheries professions, including minorities and women.

Yes!

Application Deadline: February 27, 2016

-


AFS ANNUAL MEETING

2016146th Annual Meeting

of the American Fisheries Society

August 21-25, 2016

Kansas City Welcomes You!Kansas City is an urban oasis best known for its amazing jazz music and delicious barbeque restaurants. Your confer-ence experience at the Sheraton Kansas City Crown Center promises to be a memorable one, but make sure to take the time to revel in all that KC has to offer. While in town, catch a Royals baseball game at Kauffman Stadium, discover the history of the city at the Liberty Memorial, visit the Arabia Steamboat Museum, explore hands-on exhibits inside Science City at Union Station, take in breathtaking art at the Nelson-Atkins Museum of Art, or capture the downtown night life at the Power and Light District. With plenty of Midwest charm, Kansas City has a long and diverse history for all visitors to enjoy.

Kauffman Stadium, home of the Kansas City Royals, 2015 World Series Champions. Photo credit: VisitKC.com.

View of Union Station and downtown from the Liberty Memorial. Photo credit: Missouri Division of Tourism.

Sheraton Crown Center. Photo credit: Visit KC.com.

Country Club Plaza, J. C. Nichols Fountain glowing blue in support of the Kansas City Royals. Photo credit: Will Kirby.


Association of Juvenile Salmon and Estuarine Fish with Intertidal Seagrass and Oyster Aquaculture Habitats in aNortheastPacificEstuary.Brett R. Dumbauld, Geoffrey R. Hosack, and Katelyn M. Bosley. 144:1091-1110.

[Note]FirstRecordofPugheadDeformityinBlueCatfish.Joseph D. Schmitt and Donald J. Orth. 144:1111-1116.

Exploring Crowded Trophic Niche Space in a Novel Res-ervoir Fish Assemblage: How Many is Too Many? Lisa K. Winters and Phaedra Budy. 144:1117-1128.

Spatial and Temporal Distribution of Spawning Events and Habitat Characteristics of Sacramento River Green Stur-geon.William R. Poytress, Joshua J. Gruber, Joel P. Van Eenennaam, and Mark Gard. 144:1129-1142.

A Reappraisal of Reproduction in Anadromous Alewives: Determinate versus Indeterminate Fecundity, Batch Size, andBatchNumber.Konstantinos Ganias, Jeffrey N. Divino, Katie E. Gherard, Justin P. Davis, Foivos Mouchlianitis, and Eric T. Schultz. 144:1143-1158.

Cool, Pathogen-Free Refuge Lowers Pathogen-Associated PrespawnMortalityofWillametteRiverChinookSalmon.Susan E. Benda, George P. Naughton, Christopher C. Caudill, Michael L. Kent, and Carl B. Schreck. 144:1159-1172.

Overlapping Habitat Use of Multiple Anadromous Fish SpeciesinaRestrictedCoastalWatershed.M. Chad Smith and Roger A. Rulifson. 144:1173-1183.

Effects of Water Temperature and Fish Size on Predation Vulnerability of Juvenile Humpback Chub to Rainbow Trout andBrownTrout.David L. Ward and Rylan Morton-Starner. 144:1184-1191.

InfluenceofHydrographicConditionsontheDistributionof Spiny Lobster Larvae off the West Coast of Baja Califor-nia.René Funes-Rodríguez, José A. Ruíz-Chavarríıa, Rogelio González-Armas, Reginaldo Durazo, and Sergio A. Guzmán-del Proó. 144:1192-1205.

Comparison of Two Length-Based Estimators of Total Mortality:ASimulationApproach.Amy Y. Then, John M. Hoenig, Todd Gedamke, and Jerald S. Ault. 144:1206-1219.

Effects of Spatial Extent on Modeled Relations between HabitatandAnadromousSalmonidSpawningSuccess.Steven F. Railsback, Bret C. Harvey, and Jason L. White. 144:1220-1236.

Length and Condition of Wild Chinook Salmon Smolts InfluenceAgeatMaturity.Ian A. Tattam, James R. Ruzycki, Josh L. McCormick, and Richard W. Carmichael. 144:1237-1248.

Journal HighlightsTRANSACTIONS OF THE AMERICAN FISHERIES SOCIETYVolume 144, Number 6, November 2015

InterpretingLampreyAttacksonPacificCodintheEasternBeringSea.Kevin A. Siwicke and Andrew C. Seitz. 144:1249-1262.

Trends in the Reproductive Phenology of two Great Lakes Fishes.John Lyons, Andrew L. Rypel, Paul W. Rasmussen, Thomas E. Burzynski, Bradley T. Eggold, Jared T. Myers, Tam-mie J. Paoli, and Peter B. McIntyre. 144:1263-1274.

Effects of Hybridization between Nonnative Rainbow Trout and Native Westslope Cutthroat Trout on Fitness-Related Traits.Daniel P. Drinan, Molly A. H. Webb, Kerry A. Naish, Steven T. Kalinowski, Matthew C. Boyer, Amber C. Steed, Brad-ley B. Shepard, and Clint C. Muhlfeld. 144:1275-1291.

[Comment] Comment: Natural Productivity in Steelhead Populations of Natural and Hatchery Origin—Assessing HatcherySpawnerInfluence.Richard W. Carmichael, Mark W. Chilcote, and Charles W. Huntington. 144:1292-1298.

[Comment] Natural Productivity in Steelhead Populations of Natural and Hatchery Origin—Assessing Hatchery Spawner Influence:ResponsetoComment.D. Brent Lister. 144:1298-1300.

IdentificationandDistributionofMorphologicallyCon-served Smoothhound Sharks in the Northern Gulf of Mexico.Melissa M. Giresi, R. Dean Grubbs, David S. Port-noy, William B. Driggers III, Lisa Jones, and John R. Gold. 144:1301-1310.

Development of a Bioenergetics Model for the Threespine Stickleback.Rachel A. Hovel, David A. Beauchamp, Adam G. Hansen, and Mark H. Sorel. 144:1311-1321.

Using Aerial Imagery to Characterize Redband Trout Habi-tatinaRemoteDesertLandscape.Daniel C. Dauwalter, Kurt A. Fesenmyer, and Robin Bjork. 144:1322-1339.

Patterns of Fish Assemblage Structure and Habitat Use among Main- and Side-Channel Environments in the Lower KootenaiRiver,Idaho.Carson J. Watkins, Bryan S. Stevens, Michael C. Quist, Bradley B. Shepard, and Susan C. Ireland. 144:1340-1355.

Referee Acknowledgments

End-of-Volume Author Index and Volume 144 Table of Contents


January 24–27, 201676th Midwest Fish & Wildlife Conference | Grand Rapids, Michigan | midwestfw.org

February 17–21, 2016Southern Division Spring Meeting | Wheeling, West Virginia | sdafs.org

March 2–3, 2016AFS Iowa Chapter Annual Meeting | Honey Creek Resort, Moravia, Iowa | iowa.fisheries.org

March 2–4, 2016AFS Florida Chapter Annual Meeting | Haines City, Florida | sdafs.org/flafs/meetingregistration

March 13–15, 2016Hugh C. Becker Muskie Symposium | Minnetonka, Minnesota | muskiesinc.org/anniversary

March 21–24, 2016Western Division Annual Meeting | Reno, Nevada | wdafs.org

May 21, 20162nd World Fish Migration Day | www.worldfishigrationday.com

May 23–27, 2016Planning and Executing Successful Rotenone and Antimycin Projects Course | USU, Logan, Utah | fisheries.org

May 23–27, 20167th World Fisheries Congress | Busan, South Korea | wfc2016.or.kr

June 12–16, 201612th International Congress on the Biology of Fish | San Marcos, Texas | txstate.edu/continuinged/Events/ICBF.html

June 19–23, 201640th Annual Larval Fish Conference | Solomons, Maryland | larvalfishcon.org

July 11–14, 2016Freshwater Invasives – Networking for Strategy (FINS-II) | Zagreb, Croatia | finsconference.eu

August 21–25, 2016146th Annual Meeting of the American Fisheries Society | Kansas City, Missouri | 2016.fisheries.org

August 24–25, 20163rd Annual International Conference on Fisheries and Aquaculture | Sri Lanka

September 5–8, 2016Australian Society for Fish Biology Conference | Hobart, Tasmania

October 2–6, 2016The World of Trout: 1st International Congress | Bozeman, Montana | troutcongress.org

November 10–12, 20162nd International Congress on Applies Ichthyology and Aqautic Environment | Mesolonghi, Greece | hydromedit2014.apae.uth.gr

To submit upcoming events for inclusion on the AFS website calendar, send event name, dates, city, state/ province, web address, and contact information to [email protected]. (If space is available, events will also be printed in Fisheries magazine.) More events listed at www.fisheries.org

CALENDAR


The 41st annual ASFB conference that was held in October 2015 in Sydney, New South Wales, had plenary speakers each morning covering topics as diverse as indigenous fishing rights, fish screening technology, abalone sea-ranching through to e-DNA. The meeting was held in conjunction with the International Symposium on Stock Assessment and Sea Ranching, which drew about 70 international participants of the total 300 conference attendees. This conference was large enough for there to be up to six concurrent sessions over three days, although this format varies annually. The ASFB conferences rotate geographically around the country, and New Zealand with the membership in the various states taking their turn in hosting the event, as with AFS. The next ASFB conference will be September 5-8, 2016, in Hobart, Tasmania, with Western Australia already lined up for 2017.

Here is some unique Australian fishery information from the meeting that we would like to share:• In 2011, there was an extreme marine warming event off

Southwest Australia where surface ocean temperatures were 3-5 C° above normal. This caused several tropical species to recruit and overwinter, but only the rabbitfish Siganus sp. was shown to have reproduced over 200 nm south of its historical southern limit.

• Common Carp Cyprinus carpio expansion in recent years has resulted in plans for control using a herpes virus.

• There is commercial sea ranching of abalone in Western Australia that involves the deployment of artificial reef structures that provide benefits for other marine species.

• There is increasing use of artificial reefs in marine waters for recreational fishing and a concurrent increase in research programs to monitor their success.

• The use of acoustic telemetry is very popular and large arrays of receivers are in place, particularly off Australia’s east coast.

• An innovative program called Redmap (redmap.org.au) allows citizen scientists to report observations of unusual fish and other marine life along with photographs for scientific verification.

• There are directed recreational fisheries on Atlantic Salmon Salmo salar escapees from net pen ranching in Tasmania. Escapees are not as much of a genetics issue as in North America since there are no other native salmon species.

There are many benefits of the officer exchange between AFS and ASFB. Foremost is communication of scientific information, and the meeting highlights above provide a few examples. There were several presentations at the ASFB meeting on large-scale recreational fishing surveys in Australia and New Zealand that prompted the sharing of information on new U.S. national survey methodologies. Several ASFB members gained new insights on the potential for publishing in AFS journals or attending AFS meetings. There was sharing of ideas for society governance like the Emerging Leaders Mentoring Program, involvement of women scientists in leadership positions, and student engagement strategies. Regarding students, both societies have student presentation awards and international travel awards. Perhaps there can be reciprocal study opportunities between the two societies in the future. Of note here is that Australian colleges and universities do not typically have master’s level programs like in North America.

The AFS-AFSB MOU exchange is off to a great start. This co-authored column is just the first step in mutual information dissemination. Members can attend meetings of the other society or write journal or newsletter articles. The more that happens, the more the realization that fisheries issues have commonalities, even if they are halfway around the world from each other.

Continued from p. 3BACK PAGE


AprilThis month images of Loch Ness captured by Google’s

Street View were shared. Astoundingly, large, snake-like shadows were evident in several photographs, suggesting the famed lake monster, Nessie, exists. Kidding! April Fools! Although later in the year, Nessie devotee Steve Feltham proposed the beast was a Wels Catfish Silurus glanis, introduced into UK waters in the 1860s. However, Nessie was first described as an “aquatic monster” in the book The Life of Saint Columba written in the 7th century.

MayOpa! This month celebrated the NOAA-led research

revealing that Opah Lampris guttatus have endothermic properties. This discus-shaped fish generates heat from pectoral red muscle that is activated when Opah vigorously oscillate their pectoral fins to move forward (Wegner et al. 2015). Counter-current heat exchangers in the gills curtail heat loss. These two properties facilitate circulation of warm blood throughout the Opah’s body.

JuneThis month one couldn’t help but want to pinch the cheeks,

or in this case tentacles, of the yet unnamed “flapjack” octopus. With a diameter of only 7 in (18 cm), a squishy, pink-hued body, and oversized “puppy dog eyes,” this deep-sea octopus is quite adorable. So adorable that the species may end up being named Opisthoteuthis “adorabilis.” Stephanie Bush, a postdoctoral researcher at the Monterey Bay Aquarium Research Institute (MBARI), has dedicated endless hours examining preserved and live specimens prior to formal naming. She is also patiently waiting for eggs that were unexpectedly released by a captive individual to hatch.

Year in Review: Natalie SopinkaAFS Contributing WriterE-mail: [email protected]

JanuaryThe year started off with heart-warming, yet ridiculous, news

of a dedicated goldfish owner in the United Kingdom spending around £300 (over $400 USD) on an operation to relieve his constipated pet.

February In February, we were introduced to Peckoltia greedoi,

a catfish from Brazil that bears a striking resemblance to its namesake, Greedo, the Star Wars bounty hunter pursuing Hans Solo (Armbruster et al. 2015).

The second month of the year was a historic one for Oregon Chub Oregonichthys crameri, the little fish that could. News spread quickly when Oregon Chub were taken off the federal list of endangered and threatened animals—the first fish ever to be delisted (Hughes 2015). The tremendous efforts of those involved with this recovery initiative would be recognized later in the year by AFS when the “Oregon Chub Team” was awarded the President’s Fishery Conservation Award.

MarchMarch was a tumultuous month for the Heiltsuk First Nation

in British Columbia, Canada. Conflict arose when Fisheries and Oceans Canada (DFO) approved commercial openings for the Pacific Herring Clupea pallasii roe fishery, despite the Heiltsuk forgoing their own traditional fishery to protect ravaged stocks. The DFO contended that the Pacific Herring stocks were stable enough for “modest” harvest, but the Heiltsuk were not convinced based on their assessments. Following protests and tense negotiations, the fishery was closed by DFO within a week of opening.

The catfish Peckoltia greedoi. Photo credit: Jonathan Armbruster.

BACK PAGE

2015


JulyFor Sockeye Salmon Oncorhynchus nerka in the Columbia

River, summer heat waves throughout the Pacific Northwest threatened successful migration to spawning grounds. Reduced snowpack, low water flow, and rising water temperatures were setting-off alarm bells. A similar story emerged for Sockeye Salmon migrating in Canada’s Fraser River watershed; snowmelt was meager, water levels were the lowest in 25 years, and water temperatures warmer than expected for July.

AugustThere was an enormous aggregation of over 3,000 fisheries

scientists and professionals in Portland, Oregon for the AFS Annual Meeting. The program was wide-ranging: from rockfish management, fish navigation, and science communication to skip spawning, a “Gutshop,” and film festival. Relive the meeting via Twitter by searching #AFS145.

SeptemberLast autumn, we learned of 11,500 year-old Chum Salmon

O. keta bones discovered in Alaska. The bones were found at an archeological site with a hearth located nearby a river. The find is the earliest evidence of humans in North America using this ecologically and economically valuable fish (Halffman et al. 2015).

OctoberThis month, fish were deemed the Olympians of releasing

oxygen to tissue. AFS members Jodie Rummer and Colin Brauner demonstrated that highly pH-sensitive hemoglobin in Rainbow Trout O. mykiss confer an oxygen-release advantage far superior to that of human hemoglobin (Rummer and Brauner 2015). How does this translate into underwater athleticism? When fishes put the pedal to the metal (e.g., long-distance migration, escape from a predator), delivery of much needed oxygen to exercising muscles is significantly enhanced.

NovemberNovember was Manatee Awareness Month. In the Florida

everglades, November brings the movement of the state’s of-

ficial marine mammal to warmer waters. It is estimated that ~6,000 Florida manatees Trichechus manatus latirostris reside in the waters surrounding the state (Martin et al. 2015).

DecemberThe last month of the year marked the commencement of

the ice fishing season. Across the Northern Hemisphere, fishers packed up their ice saws, chisels, rods, spears, nets, and a warm drink. Yellow Perch Perca flavescens were caught in Montreal’s Pêche Blanche Ice Fishing Village. Arctic Grayling Thymallus thymallus were caught in the lakes of Finland’s Lapland region. Carp (family Cyprinidae) were caught at the Chagan Lake Winter Fishing Festival in China. Catching fish at this historic gathering is thought to bring good fortune in the new year. Here’s to 2016!

ReferencesArmbruster, J. W., D. C. Werneke, and M. Tan. 2015. Three new species

of saddled loricariid catfishes, and a review of Hemiancistrus, Peckoltia, and allied genera (Siluriformes). ZooKeys 480:97-123.

Halffman, C. M., B. A. Potter, H. J. McKinney, B. P. Finney, A. T. Rod-rigues, D. Y. Yang, and B. M. Kemp. 2015. Early human use of anadromous salmon in North America at 11,500 y ago. Proceed-ings of the National Academy of Sciences 112(40):12344-12348.

Hughes, B. 2015. Q&A: the success story of the Oregon Chub: an interview with Paul Scheerer. Fisheries 40(8):354-355 DOI: 10.1080/03632415.2015.1065149

Martin, J., H. H. Edwards, C. J. Fonnesbeck, S. M. Koslovsky, C. W. Harmak, and T. M. Dane. 2015. Combining information for moni-toring at large spatial scales: first statewide abundance estimate of the Florida manatee. Biological Conservation 186:44-51.

Rummer, J. L., and C. J. Brauner. 2015. Root effect haemoglobins in fish may greatly enhance general oxygen delivery relative to other vertebrates. PLoS ONE 10(10):e0139477.

Wegner, N. C., O. E. Snodgrass, H. Dewar, and J. R. Hyde. 2015. Whole-body endothermy in a mesopelagic fish, the Opah, Lam-pris guttatus. Science 348(6236):786-789.

Flapjack octopus in the genus Opisthoteuthis photographed 330 m below the surface in Monterey Bay. Photo credit: 2013 MBARI.

AFS gets its name in lights. Photo credit: VEMCO.

OCEAN RIVER

CREEKLAKE

ATS has reliable aquatic tracking systems for

every environment. Live chat with a Consultant now at atstrack.com.

OCEAN RIVER

CREEKLAKE

ATS has reliable aquatic tracking systems for

every environment. Live chat with a Consultant now at atstrack.com.

Actu

al v

iew

from

the

insi

de o

f a L

arge

mou

th B

ass

Cou

rtesy

: Ric

ky D

eBol

iac

A vital question in fisheries research: what if we could definitively know when an acoustically tagged fish has been consumed by a predator? That important question is driving the evolution of HTI’s patent-pending Predation Detection Acoustic Tag (PDAT).

Designed to conclusively indicate if an acoustically tagged fish has been consumed, we’re just at the beginning of answering predation questions.

Learn more at www.HTIsonar.com/PDAT.

A Wealth of Technology & ExperienceBuilt Upon Sound Principles

(206) 633-3383www.HTIsonar.com

fisheries - amazon s3 · 38 changing philosophies of fisheries management as illustrated by the...

Documents