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Fisheries

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Download by: [Department Of Fisheries] Date: 13 March 2016, At: 23:47 FisheriesVol. 40 • No. 11 • November 2015 Downloaded by [Department Of Fisheries] at 23:47 13 March 2016

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COLUMNS PRESIDENT'S COMMENTARY 527 The Power of a Motion Ron Essig POLICY 528 Traditional Knowledge and Biodiversity Thomas E. Bigford 532 A fish biologist holds an adult male Atlantic Salmon. Photo credit: USFWS.

THE COMMUNICATION STREAM 530 Doing Cool Science? Why You Should Be Blogging about It Jeremiah Osborne-Gowey

JOURNAL REVIEWS 529 Fisheries Classics: The Model That Turned Out to Be, Sadly, but Absolutely Right Jeff Schaeffer 529 Everything You Thought about Stone Crab Fisheries Is Wrong Jeff Schaeffer BETTER KNOW A HATCHERY 532 Craig Brook National Fish Hatchery

FEATURES AND ESSAYS 534 Data Needs to Assess Effects of Soft Plastic Lure Ingestion on Fish Populations Jordan Skaggs and Micheal S. Allen 536 The Impacts of Fishing on Hermaphroditic Species and Treatment of Sex Change in Stock Assessments Mikaela M. Provost and Olaf P. Jensen 546 The Role of Introduced Populations in the Management and Conservation of Least Chub Rosebud Pond, an example of a habitat where Least Chub were Paul D. Thompson, P. Aaron Webber, and Cassie D. Mellon 546 introduced. Photo credit: Paul Thompson. 557 Relevant Topics to Keep in Mind as a Graduate Student Artur Rombenso, Karma Kissinger, Michael Ciaramella, and Jesse Trushenski

562 Enhancing the Utility of the NHDPlus River Coverage: Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 Characterizing Ecological River Reaches for Improved Management and Summary of Information Daniel J. Wieferich, Wesley M. Daniel, and Dana M. Infante

Barracuda - say cheese! Photo credit: Florida Fish and Wildlife 571 Conservation Commission.

Fisheries | www.fisheries.org 525 565 146TH ANNUAL MEETING OF THE AMERICAN FISHERIES SOCIETY: THIRD CALL FOR PAPERS Fisheries JOURNAL HIGHLIGHTS American Fisheries Society • www.fisheries.org 566 North American Journal of Aquaculture Volume 77, Number 4, October 2015 EDITORIAL / SUBSCRIPTION / CIRCULATION OFFICES 5410 Grosvenor Lane, Suite 110•Bethesda, MD 20814-2199 568 CALENDAR (301) 897-8616 • fax (301) 897-8096 • [email protected] The American Fisheries Society (AFS), founded in 1870, is the 568 CORRECTION oldest and largest professional society representing fisheries scientists. The AFS promotes scientific research and enlight- ened management of aquatic resources for optimum use and BACK PAGE enjoyment by the public. It also encourages comprehensive 571 The Teeth of Fishes: Say Ahhh! education of fisheries scientists and continuing on-the-job Natalie Sopinka training.

AFS OFFICERS EDITORS PRESIDENT CHIEF SCIENCE EDITORS Ron Essig Jeff Schaeffer Olaf P. Jensen PRESIDENT-ELECT Joe Margraf SCIENCE EDITORS Kristen Anstead FIRST VICE PRESIDENT Marilyn “Guppy” Blair Steve L. McMullin Jim Bowker Mason Bryant SECOND VICE PRESIDENT Steven R. Chipps Jesse Trushenski Ken Currens Andy Danylchuk PAST PRESIDENT Michael R. Donaldson Donna L. Parrish Andrew H. Fayram Stephen Fried EXECUTIVE DIRECTOR Larry M. Gigliotti Doug Austen Madeleine Hall-Arbor Alf Haukenes FISHERIES STAFF Jeffrey E. Hill SENIOR EDITOR Deirdre M. Kimball Doug Austen Jeff Koch Jim Long DIRECTOR OF Daniel McGarvey PUBLICATIONS Jeremy Pritt Aaron Lerner Roar Sandodden Jesse Trushenski MANAGING EDITOR Usha Varanasi Sarah Harrison Jeffrey Williams

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 BOOK REVIEW EDITOR CONTRIBUTING EDITOR Francis Juanes Sarah Fox COVER ABSTRACT TRANSLATION CONTRIBUTING EDITOR Pablo del Monte-Luna Beth Beard ARCHIVE EDITOR Mature male Black Sea Bass displaying secondary sex Mohammed Hossain characteristics (bright blue coloration and forehead hump) CONTRIBUTING WRITER typical of males during the spawning season. Photo credit: Natalie Sopinka Olaf P. Jensen.

Fisheries (ISSN 0363-2415) is published monthly by the American DUES AND FEES FOR 2015 ARE: Fisheries Society; 5410 Grosvenor Lane, Suite 110; Bethesda, MD 20814-2199 © copyright 2015. Periodicals postage paid at Bethesda, Maryland, and $80 in North America ($95 elsewhere) for regular members, $20 in at an additional mailing office. A copy of Fisheries Guide for Authors is North America ($30 elsewhere) for student members, and $40 ($50 available from the editor or the AFS website, www.fisheries.org. If request- elsewhere) for retired members. ing from the managing editor, please enclose a stamped, self-addressed envelope with your request. Republication or systematic or multiple repro- Fees include $19 for Fisheries subscription. duction of material in this publication is permitted only under consent or license from the American Fisheries Society. Nonmember and library subscription rates are $191. Postmaster: Send address changes to Fisheries, American Fisheries Society; 5410 Grosvenor Lane, Suite 110; Bethesda, MD 20814-2199.

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526 Fisheries | Vol. 40 • No. 11 • November 2015 COLUMN PRESIDENT'S COMMENTARY

The Power of a Motion Ron Essig, AFS President AFS President Ron Essig [email protected]

We’ve probably all experienced the following scenario. Sometimes the group struggles to come to a consensus on An idea surfaces during lengthy discussion in a meeting that wording of a motion. This is when a chair can take advantage could move a group forward. Several meeting participants agree of a meeting break to allow time for a couple members to work that the idea is a good one. With luck, this idea is recorded in on wording that captures the flavor of the points raised. After meeting minutes; however, it is never acted on later. With typical the break, one of those individuals offers the motion. Hopefully, member turnover in any group, it may be months or years before there is a second to the motion, and the ensuing discussion might the issue resurfaces. Then members struggle to recall that good be more straightforward. There can be many permutations to the idea and why it was not implemented. simplified process described here, but it is not my intent, nor my What can prevent this scenario from occurring is taking expertise, to discuss them here. action at the time an idea surfaces. For an informal group, this might simply be the leader sensing buy-in from most members An effective strategy with motions is to and proceeding in that direction. This is a typical process in many AFS committees, which have relatively small numbers of get support behind it before you introduce members. However, in larger AFS Units—Sections, Divisions, and Chapters—there needs to be a bit more formality to maintain it. This means doing some homework an orderly process. I am a poor constitutional consultant, but I vetting it prior to the meeting. have learned that the motion process within Robert’s Rules of Order is a very powerful tool indeed. It is what moves forward You don’t want your idea to fall on deaf the Society, its Units, and other organizations that you may be part of in your work and personal life. ears and go nowhere. According to Robert’s Rules, a motion is simply a formal proposal by a member that the group take a certain action. The AFS is fortunate to have the appointed position of process involves someone making a motion for action that is constitutional consultant. Each new person taking this on has germane to the particular agenda topic. That person should state a one-year apprenticeship with their predecessor, so he/she is the exact wording of the motion. It can be modified through well-trained and hits the job running. This individual is well subsequent dialogue with the chair of the meeting. Then if versed in Robert’s Rules and the AFS Constitution, Rules, and modified, the chair should restate the motion and obtain a second Procedures. They also have to qualify for membership in the to the motion before it can be debated. The purpose of a second National Association of Parliamentarians. They draw on that is to indicate that more than one member of the group agrees knowledge base to offer constructive criticism of draft motions that the motion should be discussed. The seconder does not have on a variety of AFS issues, including Unit bylaw changes. It is to agree with the motion. After discussion, either a vote is taken particularly important for the constitutional consultant to review Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 or the motion is declared passed by unanimous consent. draft motions to come before the Management Committee or the An effective strategy with motions is to get support behind Governing Board. Getting clear wording will save considerable it before you introduce it. This means doing some homework time debating what the desired purpose of the motion really is. vetting it prior to the meeting. You don’t want your idea to fall There have been several examples in my officer tenure when on deaf ears and go nowhere. Having at least a couple members motions have been sent back for reworking, which could have on your side may help sway the conversation to the conclusion been avoided with constitutional consultant input beforehand. you want. Depending on complexity, it is desirable to write I hope these words encourage you to offer motions during out proposed motions either before the meeting or during the business meetings of your AFS Unit. If you don’t have the meeting. That way, there is no uncertainty in exactly what is wording just right, it is likely that someone else will offer being requested since misstating a key word can make a big amended language that will work better for the group. The difference. power of a motion can only be realized if it is presented to the group. Take action! Make motions!

Fisheries | www.fisheries.org 527 COLUMN POLICY Traditional Knowledge

and Biodiversity AFS Policy Director Thomas E. Bigford, AFS Policy Director Thomas E. Bigford [email protected]

This month’s column is about the intersection of human of nearshore marine species were probably healthier centuries and biological diversity with history. According to McGinley ago when fewer river blockages hampered migrations of (2014), “[s]pecies diversity is a measure of the diversity within preferred prey. While we have difficulty modelling predator an ecological community that incorporates both species richness (tuna and Striped Bass Morone saxatilis) and prey (river (the number of species in a community) and the evenness herring and menhaden) populations back to colonial times, we of species’ abundances.” Beyond biology, and bridging the might gain insights by investigating Tribal and early colonial social and natural sciences, traditional knowledge is gathered histories. Perhaps Indian middens will reveal the historical from people with diverse yet intimate perspectives—historical range of anadromous fish and the health of marine predators that observations by those close to a situation, cultural and economic benefitted from stronger food webs. Even reviewing colonial aspects, and other personal insights—that provide a solid basis newspapers would be enlightening, as done by John McPhee for resource management decisions and policies. (2002) in The Founding Fish. A stronger mix of natural and social sciences should yield Traditional knowledge is gathered from benefits to fisheries professionals and to our efforts to manage fish. History informs the present and helps us to anticipate people with diverse yet intimate perspectives the future. I see an interesting parallel to efforts decades ago that provide a solid basis for resource to design more inclusive environmental decision processes, but now with an emphasis on knowledge rather than political management decisions and policies. persuasion or ethnicities. So how should we proceed? How do we add valuable These thoughts were inspired by the AFS 2015 Annual perspectives to our work in the sciences, fishery management, Meeting. Amid thousands of presentations on topics from the and policy? First, you need to challenge my assumptions and our Mekong River to fatty acid signatures, my personal eureka current efforts to add traditional knowledge. Are we doing better moment was a talk by Dionne Hoskins of NOAA Fisheries (also in some topical areas? Are there best practices in gathering professor at Savannah State University; speaking at the Equal or sharing knowledge? Second, after we focus on needs, we Opportunities Section meeting). Hoskins described “Voices must enlist experts to increase our prospects for success. That from the Fisheries,” a NOAA Fisheries (2015) repository reach should reflect the ethnic and cultural diversity of those for “consolidating, archiving, and disseminating oral history experts. Jeremy Pyatskowit (past president, AFS Native Peoples interviews related to commercial, recreational, and subsistence Fisheries Section, and member of the Menominee Indian Tribe fishing. . . .” Oral histories are a treasure trove of human of Wisconsin) recommended we seek this knowledge by inviting dimension insights that can help to expand our horizons and under-represented voices into AFS business across all Units improve success. During her talk, I recalled comparable efforts and activities. Adding diversity to all discussions will lead to Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 in Maine to gather local histories from the lobster industry, greater results than we would see from a separate discussion on centuries of perspective available from Native Peoples on every traditional knowledge. He even suggested AFS forego efforts continent, and other efforts. to reinvigorate its nascent Native Peoples Fisheries Section and Hoskins’ talk affirmed that a significant slice of our history instead add those issues to our expectations of every AFS Unit. is not recorded in journals and proceedings. This is frustrating Finally, we need to appreciate the body of knowledge awaiting but not very surprising. After we struggle to locate those with inclusion. I strongly suspect there are efforts around our valuable insights, the lessons to learn are not always preserved waterways to gather local knowledge. Let’s use Dionne Hoskins’ in typical formats. We’re doing better with social media and work to inspire us. audio-video recordings, which capture content sometimes lost in With an eye toward historical, cultural, and topical printed materials. This is crucial, as that traditional knowledge— inclusiveness across all of our work, I suspect we will all be those historical voices reflecting cultural history—offers us an more successful. opportunity to develop more successful management strategies and policies. REFERENCES The real opportunity lies with collaboration, i.e., merging McGinley, M. 2014. Species diversity. Encyclopedia of Earth. Avail- local knowledge with the social and natural sciences, resource able: www.eoearth.org/view/article/156211. (September 2015). McPhee, J. 2002. The founding fish. Farrar, Straus, and Giroux. New management, policy, etc. I can imagine how traditional insights York. could help us to resolve resource management challenges that, NOAA Fisheries. 2015. Voices from the fisheries. NOAA. Available: in the absence of strong data sets, tend to be more qualitative www.st.nmfs.noaa.gov/humandimensions/voices-from-the- than quantitative. For example, logic dictates that populations fisheries/index. (September 2015).

528 Fisheries | Vol. 40 • No. 11 • November 2015 JOURNAL REVIEWS Fisheries Classics: The Model That Turned Out to Be, Sadly, but Absolutely Right Jeff Schaeffer | AFS Co-Chief Science Editor. E-mail: [email protected]

The upper Laurentian Great Lakes were separated physically from the Atlantic coast by Niagara Falls, but constructed ship canals allowed marine invaders to penetrate the system. Invading Sea Lampreys Petromyzon marinus decimated native piscivores during the 1940s, which allowed Alewife Alosa pseudoharengus to proliferate to nuisance levels of abundance by the 1960s. Fisheries managers attacked the Alewife problem by introducing Pacific salmonOncorhynchus spp. This worked, and created a wildly popular sport fishery that had not existed previously. This was a classic “be careful what you wish for, because you may get it” conundrum. There was some evidence that Alewives were being controlled, and there was tremendous stakeholder pressure to maximize salmon populations because the novel sport fishery was actually revitalizing coastal communities. Indeed, Lakes Michigan and Huron resembled a cornucopia in that there was no appar- ent upper limit in terms of how many predators could be sustained. However, in 1981, Don Stewart, Jim Kitchell, and Larry Crowder published a Transactions paper that suggested there was an upper limit to how many predators could be sustained. Using bioenergetics modeling (a rather novel concept at that time), they suggested that stocking programs had decoupled predatory demand from prey production, with possible destabilization of the predator-prey system and ultimately strong potential effects on Alewife biomass, with most consumption by Chinook Salmon O. tschawytsha. Their work created a firestorm in that it was the basis for a wide range of subsequent applications of their method to evaluate other predator-prey systems (many of those manuscripts appeared in TAFS), but in the Great Lakes it began a controversy that raged for the next 25 years. There were some managers and researchers who began calling for more judicious stocking rates, while others expressed utter disbelief that a stocking program could actually impact prey dynamics within one of the largest freshwater ecosystems on the planet. No real consensus emerged, but a regional approach developed in which both predators and prey were assessed annually, stocking rates were reduced somewhat from initial levels, and predator-prey dynamics were studied intensively. This approach sustained the salmon fishery until 2004 when Alewife collapsed in Lake Huron, with concurrent decline in Lake Michigan. While there is still some debate regarding the role of climate and invasive effects as contributing factors, salmonid predation was implicated strongly as being severe enough to serve as an absolute slate-wiper for Alewife, with both stocked and feral fish contributing to a high predatory demand. And while salmon controlled a malignant invader, there was an economic burden placed on coastal communities, especially in Lake Huron where fishing effort declined sharply in many areas. On the other hand, Alewife absence was followed by strong recruitment of several native fishes that had experienced chronic low abundance for decades in the presence ofAlewife. We love this paper because it was so prescient, and it stimulated a broad range of new research efforts that led to great insight into how models can be useful in understanding predator-prey dynamics. And it began a compelling story about a bold management strategy that led to initial great success, but eventually experienced the most feared and unlikely outcome. REFERENCE Stewart, D. J., J. F. Kitchell, and L. B. Crowder. 1981. Forage fishes and their salmonid predators in Lake Michigan. Transactions of the American Fisheries Society 110(6):751-763. dx.doi.org/10.1577/1548-8659(1981)110<751:FFATSP>2.0.CO;2 Everything You Thought about Stone Crab Fisheries Is Wrong The Stone Crab Menippe spp. fishery is one of the most unusual fisheries in the world, because only claws are harvested, and animals are released to remain in the fishery via survival and claw regeneration. Florida accounts for the majority of landings, and

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 there both claws can be removed if they are of legal size, whereas in South Carolina, only one legal-sized claw can be removed from a single crab. One of the major drivers of this fishery is the assumption that stone crabs survive claw harvest and regenerate, but Elizabeth Duermit (South Carolina Department of Natural Resources, Marine Resources Research Institute and Grice Marine Laboratory, College of Charleston) and her colleages found that this may not be the case due to both direct and indirect effects. They used a series of elegant experiments to determine that wound widths greater than 7 mm caused high post-removal mortality, and that wound size was more important than if one or both claws were taken. This suggests that fisher skill with claw removal was an important component underlying short-term survival, but there were also long-term effects in that stone crabs with one claw consumed fewer bivalves, and stone crabs with no claws could consume only fish flesh. Restricted feeding could exacerbate an already-long intermolt claw regeneration interval such that few stone crabs return to the fishery. And by examining stone crab claws for sale in retail environments, they found that 20% of the claws exhibited breakage consistent with high short-term mortality. We featured this paper because it was elegant and creative, but also because it is a great example of how some long-held beliefs do not hold up when actually tested. And their work suggests that at least a proportion of mortality depends on fisher skill. The Florida Fish and Wildlife Commission (FFWC) has already established video outreach efforts to teach claw removal (youtube.com/ watch?v=YTgXTS8gLjU), but new knowledge of long-term effects may need to be considered to maintain this fishery, as well as outreach directed at providing science education to Youtube commenters. REFERENCE Duermit, E., P. R. Kingsley-Smith, and D. H. Wilber. 2015. The consequences of claw removal on stone crabs Menippe spp. and the ecological and fishery implications. North American Journal of Fisheries Management 35(5):895-905. dx.doi.org/10.1080/02755947.2015.1064836

Fisheries | www.fisheries.org 529 COLUMN THE COMMUNICATION STREAM Doing Cool Science? Why You Should Be Blogging about It Jeremiah Osborne-Gowey Jeremiah Osborne-Gowey, Publishing research in journals certainly has an esteemed and aware–perhaps painfully so–that our AFS Social Media Guru valuable place in the scientific process, and for many good reasons. writing skills have been honed on E-mail: jeremiahosbornegowey@ gmail.com But have you ever wanted to see your work reach a broader audience the academic whetstone and geared Twitter: @JeremiahOsGo and have a bigger impact beyond traditional academic boundaries? heavily toward a highly technical and Blogging may just be your answer. While microblogging platforms ever smaller audience. But in this age like Twitter, Facebook, LinkedIn, and Google Plus–along with a host of increased scrutiny of science, dwindling science budgets and rapid of others–allow for the distribution or exchange of short messages and far-reaching electronic communication, demonstrating the relevance and play a key role in communicating today, blogs are also important of science to the "bigger picture" arguably ought to be a key component communication platforms that allow for far richer and more fully of any new science undertaking. Writing for general audiences presents developed communiqués. a host of challenges to researchers most familiar with academic writing. Those challenges (e.g., finding a good hook/tying it to the human story, What are blogs? In the simplest sense, blogs are an online setting up the characters, identifying conflict, etc.) are surmountable. discussion or a listing of discussions. They originated in the late 1990s The more you practice writing, meaning the more content you produce, as daily web logs (later shortened to blogs) but gained traction later that the better your writing skills become. Blogging is an excellent platform decade and early in the 2000s with the proliferation of web publishing to help sharpen your writing skills. tools. Blogging became mainstream during the 2002 and 2004 political campaigns in the United States as pundits and news organizations widely Get to know (and build) your community. While adopted blogging as a means of rapidly and effectively communicating exact statistics are tough to come by (Blogger, the largest blogging stories. Today, there are tens of millions of visitors to blogs each month, platform, keeps their stats under wraps), best estimates put the and blogs now rival mainstream media sites for primary sources of news number of public blogs well above 300 million. Blogs are typically and information. characterized/organized into groups or communities, making it User statistics aside, there are a multitude of very good reasons for increasingly easy to find others that share similar interests. Finding out scientists to get involved in blogging. While space and time prevent me the interests of your community will help you better frame your science from covering all of them, here are the top six reasons why I believe and may lead to greater public support for it. Given nearly one quarter scientists should add blogging to their communication portfolios: of total internet usage time is devoted to social networks and blogs and nearly >75% of internet users read blogs, blogging will help you Share your expertise. Though there is some variation by connect with new and interesting people (including other scientists). topic and origination point, survey results generally indicate the public Additionally, interacting with others has repeatedly been shown to by-and-large moderately to highly trust scientists, especially those from help people build their networks. The comment section of your blog is academic institutions. And the public wants to hear from scientists a fantastic way to interact with your audience, increase recognition of themselves, NOT from politicians that spin the science. What’s more, you or your organization or about a particular issue, and help grow your the public is less and less trusting that major news outlets are reporting exposure and audience. unbiased stories while public trust in blogs, especially individual blogs and blogs not backed by governments, interest groups or corporations, Increase your "findability" in search engines. is on the rise. There is a relatively small but growing body of research People need to find your blog to read it and the "finding" is driven examining the tangible impacts of science blogging but if (1) scientists by search engines. Google, Yahoo, Bing, and others keep their search want to get research findings out to a broader audience, (2) the public algorithms under tight wraps. Over the years, however, research on trusts scientists and wants to hear directly from them, and (3) the public search engine optimization (SEO, for short) has provided clues for how

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 generally trusts information and advice from individual blogs, it is not a to make you and your organization more "findable" in Internet searches big leap to say that scientists need to be blogging. Not surprisingly, some (e.g., show up higher in the search results list). One of the best ways for of the most read blogs are either science-related (e.g., NASA, Discovery, search engines to find you or your organization is via blogging. Research Wired Science, HowStuffWorks, Pharyngula) or have a prominent indicates that search engines are much more likely to list you or your science section (e.g., Huffington Post, Boing Boing, TechCrunch, organization higher in the results from a web search if your websites Mashable, etc.). The public wants to know what you, dear scientist, have include blogs. What’s more, blogs that include keywords and links to to say. other web addresses in their posts have even more user traffic at their sites from search engine results. In short, if you want to raise the profile Personalize your science. It’s no secret that the way of your organization or about a particular topic, blog about it. we communicate is in the midst of a major paradigm shift. No longer Check out Wiki-How and HowToStartABlog, for some simple is print media king. No longer are major news organizations the considerations and step-by-step instructions. For a thorough treatment gatekeepers of what stories are fit for public consumption. Journalists of science blogging, check out and The Guardian’s recent article or are certainly not the only ones interpreting and telling the stories. True, Bora Zivkovic’s excellent Scientific American blog post from last year. some scientists have a steep learning curve when it comes to telling Regardless of which paths you take on your journey to better science compelling stories (i.e., finding the "hook") to the general public. But communication or whether you are reluctant to take it on, blogging is scientists are increasingly being sought out to tell the stories of their own one of the most effective ways of personalizing your science, honing research. For good or bad, scientists can now highlight the findings and your writing skills, raising awareness about your organization’s work or drawbacks most important in their research to minimize misconceptions, a particular topic, building a community, and increasing the likelihood misinterpretations, and inaccuracies. that internet users find your information. Despite the learning curve of moving between writing for academics and the general public, that Improve your writing skills. Refining a craft takes sounds like a winning combination to me. practice. Lots of practice. Writing is no different. Scientists are typically

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Fisheries | www.fisheries.org 531 BETTER KNOW A HATCHERY Craig Brook National Fish Hatchery What is the name of your hatchery, how did it get that name, and how long has it been in operation? Craig Brook National Fish Hatchery (NFH), in East Orland, Maine, is named after Craig Pond, which supplies the hatchery with its water. The hatchery was established in 1889 by Charles G. Atkins, who served as its superintendent until 1914 when he was appointed fish culturist-at-large. He officially retired in August 1920. What fish do you raise, approximately how many do you raise, and what are they used for? Craig Brook NFH works with partners for the conservation and recovery of Atlantic Salmon Salmo salar in the Gulf of Maine Distinct Population Segment. We culture Atlantic Salmon in an effort to prevent its extinction and preserve the genetic diversity of the populations. We typically have up to 6,000 brood fish on hand at the hatchery at any given time. We produce approximately 5 million eggs annually, which are stocked at various life stages throughout Maine waters. We supply eggs for fry stocking to Green Lake NFH and the non-governmental organization Downeast Salmon Federation (DSF) hatcheries. How big is your hatchery? Craig Brook NFH is a state-of-the-art facility, which sits on more than 130 acres of land, constructed with enough isolation and biosecurity to be seven separate hatcheries in one. We have more than 40 rearing tanks in a variety of sizes, which equates to over 34,000 cubic feet of rearing space for fish. We have 1,800 cubic feet of rearing space in 225 start-up troughs for fry and 1,150 heath incubator trays, which allows for incubation of over 9 million salmon eggs. Also located at the hatchery are two public use areas, Alamoosook Lake and Craig Pond, with boat ramps and swimming areas, as well as several miles of nature trails. Additionally, we have both an effluent and an influent wastewater treatment plant. What is the biggest challenge facing your hatchery today? What challenges do you foresee in the future? One of the biggest challenges facing the work and mission of our hatchery today is marine survival of Atlantic Salmon. Salmon are an anadromous species and therefore spend a large portion of their life in ocean waters. There are still many unknowns and unanswered questions when it comes to their survival at sea, making it a challenge for us to develop solutions. Some challenges we foresee into the future, and are still learning about, are the effects of climate change on their survival. Innovation is a part of how any operation deals with emerging challenges. How does innovation happen at your facility and how does it benefit your operation and others? Innovation is a part of dealing with emerging challenges. We work cooperatively with many partners, keeping up-to-date on new research, technologies, and scientific developments. Working with others has led to enhanced efforts in facility operations and the work we do for salmon recovery. Any recent successes, news, trivia, or facts you can share? We have been involved with many changes over time in the way salmon are cultured, spawned, and stocked to increase their survival. Each year we provide fish to scientific research for a variety of Craig Brook National Fish Hatchery located in East Orland, Maine. The hatchery celebrated its 125th anniversary this past studies. Last year, Craig Brook NFH celebrated its 125th anniversary. Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 year. Photo credit: USFWS. Are you and/or the staff at your facility active in AFS? Some Craig Brook NFH staff are active in AFS, which always provides benefits and opportunities for consultation with other fisheries professionals working in fisheries sciences. In one sentence, why is fish culture important? Fish culture is important in our efforts to help prevent extinction and the loss of genetic diversity in fish populations.

We thank Chris Domina, hatchery manager, and Peter Lamothe, complex manager, at the Craig Brook NFH for answering our questions and providing photos. To see the complete “Better Know a Hatchery” feature for Craig Brook NFH, as well as other featured facilities, visit the AFS Fish Culture Section website at: fishculturesection.org and click on the “Better Know a Hatchery” tab. Also, visit us on Facebook to see photos from all of the facilities A fish biologist holds an adult male Atlantic Salmon. Photo credit: USFWS. featured in “Better Know a Hatchery.”

532 Fisheries | Vol. 40 • No. 11 • November 2015 A fish biologist sorts through Atlantic Salmon eggs at the A group of brood stock Atlantic Salmon. Photo credit: USFWS. hatchery. Photo credit: USFWS.

Brood stock holding pools at the hatchery. Photo credit: Entrance to the visitor center at Craig Brook National Fish USFWS. Hatchery. Photo credit: USFWS. Downloaded by [Department Of Fisheries] at 23:47 13 March 2016

Fisheries | www.fisheries.org 533 ESSAY

Data Needs to Assess Effects of Soft Plastic Lure Ingestion on Fish Populations Jordan Skaggs Fisheries and Aquatic Sciences, University of Florida, Gainesville, FL

Micheal S. Allen Fisheries and Aquatic Sciences, University of Florida, 7922 NW 71st St., Gainesville, FL 32653. E-mail: [email protected]

Soft plastic lures (SPLs) are among the most common with sample sizes >500 fish (Trippel et al. 2015; Wes Porak, tackle components used in recreational fisheries worldwide. Florida Fish and Wildlife Conservation Commission, personal Local and national media sources recently raised concern communication). Biologists often attributed a category as “other, about the effect of discarded SPLs on fish populations (Raison miscellaneous, inorganic material, or unidentifiable” (Schramm et al. 2014). Fish may consume SPLs directly from baits or and Maceina 1986; Cortes 1997; Wheeler and Allen 2003; after being discarded by anglers into waters. However, it is Sammons and Maceina 2006), and it was unclear whether this uncertain whether SPLs cause individual or population-level grouping included any SPLs. Although formal documentation effects upon fish growth and survival. We reviewed existing of SPL ingestion by inland sport fishes was rarely observed in literature regarding individual and population-level effects from published diet studies, this may be because biologists rarely discarded SPLs using the key words “soft plastic lure ingestion,” record SPL ingestion events during field studies. Thus, there is “soft plastic lure consumption,” “fish diet,” and “fish stomach inconsistency in diet reporting, and the degree to which SPLs content” in the Google Scholar search engine. Our goal was to are ingested across fish species remains unknown. Given the assess the degree to which soft plastic lures have been consumed increased recent concern among biologists and legislators, there by fish and identify any individual or population-level effects. is a clear need to monitor the occurrence of SPL ingestion for a Results of this study can inform future data collection efforts in range of recreational fisheries. Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 order to test for the effects of SPLs on individual fish and fish Studies testing for the effects of SPL ingestion on fish populations. growth and survival are also rare. Danner et al. (2009) showed Stomach content analysis represents the primary method the potential consequences of SPL ingestion in the laboratory for detecting SPL ingestion in wild fish. Fish diet studies are using juvenile Brook Trout Salvelinus fontenalis (mean ± SD common and provide an understanding of factors affecting length of 234 ± 20 mm TL). An established, pellet-based feeding fish growth and survival (Hyslop 1980; Bowen 1996). Diet regime was later supplemented with various forms of SPLs studies typically categorize food items to a range of taxonomic commonly used by recreational anglers. Sixty-three percent of levels (Bowen 1996). Analyses are typically comprised of the trout (n = 37) ingested at least one SPL, and consumption of counts, frequency of prey item occurrence, volume or weight SPLs caused reductions in body weight and condition (Danner et of individual prey items, and prey energy density (Hynes 1950; al. 2009). However, the experiment’s feeding design could have George and Hadley 1979; Hyslop 1980; Hodgson and Kitchell caused conditioning effects to inflate ingestion rates of SPLs 1987). relative to conditions expected in the wild. For example, feeding Studies that report the occurrence or abundance of SPL pellets to hatchery Brook Trout and Atlantic Salmon Salmo ingestion by wild fish populations are rare. Outside of a select salar produced surface-oriented feeding behavior, and fish were few studies (i.e., Danner et al. 2009; Raison et al. 2014), more likely to consume objects dropped from above than natural common dietary data collection reports have not characterized forage items (Mason et al. 1967; Sosiak 1978). Thus, there is a the occurrence of SPL ingestion. For example, analyses of need to test for effects of SPL ingestion on growth and mortality Largemouth Bass Micropterus salmoides diets in Florida under natural feeding conditions. suggested that the occurrence of SPLs in stomachs was <1%,

534 Fisheries | Vol. 40 • No. 11 • November 2015 Danner et al. (2009) clearly showed the potential for SPLs Danner, G. R., J. Chacko, and F. Brautigam. 2009. Voluntary ingestion to influence growth rates in Brook Trout, which ultimately could of soft plastic lures affects Brook Trout growth in the laboratory. North American Journal of Fisheries Management 29:352-360. influence fisheries. Similar laboratory studies are needed for George, E. L., and W. F. Hadley. 1979. Food and habitat portioning other fish species. Raison et al. (2014) quantified potential SPL between Rock Bass (Ambloplites rupestris) and Smallmouth effects on fish populations through a combination of laboratory Bass (Micropterus dolomieu) young of the year. Transactions of the American Fisheries Society 108:253-261. experiments and field sampling (snorkel surveys, creel surveys, Hodgson, J. R., and J. F. Kitchell. 1987. Opportunistic foraging by and stomach content analysis) at Charleston Lake, Ontario. Largemouth Bass (Micropterus salmoides). American Midland Snorkel surveys found SPL presence at rates ranging from 0 Naturalist 118:323-336. to 8 SPLs per 100 m of shoreline (mean = 3.25, Raison et al. Hyslop, E. J. 1980. Stomach contents analysis: a review of methods and their application. Journal of Fish Biology 17:411-429. 2014). During the study’s creel survey, 25 of 42 anglers (60%) Hynes, H. B. N. 1950. The food of freshwater sticklebacks (Gaster- who kept Lake Trout S. namaycush reported finding SPLs in osteus aculeatus and Pygosteus pungitius) with a review of Lake Trout stomachs. Ten of the 42 anglers who harvested Lake methods used in studies of the food of fishes. Journal of Animal Ecology 19:36-58. Trout (24%) indicated that SPLs were found in over 50% of MDIFW (Maine Department of Inland Fisheries and Wildlife). 2014. fish stomachs (Raison et al. 2014). However, stomach content Report back to legislature on resolve chapter 18. Available: analysis of fish from gill-net sampling and hook-and-line www.maine.gov/ifw/pdfs/FINAL%20Report%20to%20the%20 angling revealed that only 2.2% of Lake Trout stomachs (n = Joint%20Standing%20Committee%20on%20Inland%20Fisher- ies%20and%20Wildlife%20CT%20AE%20DD%20edits12-13-13. 90 total stomachs observed) and 3.4% of Smallmouth Bass M. pdf. (September 2015). dolomieu stomachs (n = 88 total stomachs observed) contained Mason, J. W., O. M. Brynilson, and P.E. Degurse. 1967. Comparative SPLs. Thus, much uncertainty remains about the degree to survival of wild and domestic strains of Brook Trout in streams. Transactions of the American Fisheries Society 96:313-319. which fish ingest SPLs, and future work should utilize both Raison, T., A. Nagrodski, C. D. Suski, and S. J. Cooke. 2014. Exploring angler reports and scientific collections to assess ingestion rates. the potential effects of lost or discarded soft plastic fishing lures To date, no studies have indicated any individual or on fish and the environment. Water Soil Air Pollution 225:1869. population-level effects of SPLs on fisheries, but one state Sammons, S. M., and M. J. Maceina. 2006. Changes in diet and food consumption of Largemouth Bass following large-scale Hydrilla legislature has considered a ban on SPLs (MDIFW 2014). reduction in Lake Seminole, Georgia. Hydrobiologia 560:109- Our review revealed little information about ingestion rates 120. or impacts of SPL consumption on fish and fisheries. Thus, Schramm, H. J., and M. J. Maceina. 1986. Distribution and diet of Su- wannee Bass and Largemouth Bass in the lower Santa Fe River, regulations to restrict use of SPLs in order to protect fish Florida. Environmental Biology of Fishes 15:221-228. populations and fisheries would not currently be based on Sosiak, A. J. 1978. The comparative behavior of wild and hatchery- scientific proof of impacts. This issue warrants additional reared juvenile Atlantic Salmon (Salmo salar L.) Master’s thesis, experimental studies in natural and controlled environments to University of New Brunswick. Fredrickton, Canada. Trippel, N. A., M. S. Allen, and R. S. McBride. 2015. Importance of resi- test for effects of SPL ingestion on fish behavior, growth, and dent and seasonally transient prey to Largemouth Bass in the survival. Uncertainty remains about SPL residence time in the St. Johns River, Florida. Transactions of the American Fisheries environment (e.g., Raison et al. 2014). Additionally, diet studies Society 144:140-149. Wheeler, A. P., and M. S. Allen. 2003. Habitat and diet partitioning should include SPLs as a category to quantify ingestion rates. between Shoal Bass and Largemouth Bass in the Chipola River, Angler education programs emphasizing proper SPL Florida. Transactions of the American Fisheries Society 132:438- disposal and heightened industry awareness among SPL 449. manufacturing companies may serve as an alternative to SPL restrictions (Raison et al. 2014). Some fishing tackle industry and angler-driven initiatives have sought to reduce discarding of SPLs (pledgetopitchit.org) and promoted recycling (rebaits.org). The manufacturers of SPLs recognize potential environmental issues of SPL use and offer biodegradable alternatives A GRADER THAT MAKES (recycledfish.org/safe-angling/product-profiles/berkley-gulp. htm). However, metrics used by manufacturers to label SPL Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 YOUR JOB EASIER products as biodegradable are unknown (Raison et al. 2014). Now you can adjust grading sizes without juggling Thus, there is a clear need for better data on the occurrence multiple baskets. Our graders float, adjust quickly rate of SPL ingestion by fish in the wild as well as tests for any and have larger bars so fish pass through with little individual and population-level effects. We hope this essay can chance of gill damage. Perfect for fresh or salt water, help guide future studies to evaluate the effects of SPL ingestion choose either the 30 or 50 settings model. on fish populations and fisheries. 15” x 17” x 11.5” Check out our aerators ACKNOWLEDGMENTS Larger size also available. and feeder too! We would like to thank Steven J. Cooke and Gene Gilliland for comments that improved a draft of this manuscript.

REFERENCES Bowen, S. 1996. Quantitative description of the diet. Pages 513–532 Visit www.freshflo.com in B. R. Murphy and D. W. Willis, editors. Fisheries techniques. for more details. American Fisheries Society, Bethesda, Maryland. Cortés, E. 1997. A critical review of methods of studying fish feeding Wisconsin • USA • 920-208-1500 based on analysis of stomach contents: application to elasmo- branch fishes. Canadian Journal of Fisheries and Aquatic Sci- ences 54:726–738.

Fisheries | www.fisheries.org 535 FEATURE The Impacts of Fishing on Hermaphroditic Species and Treatment of

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 Sex Change in Stock Assessments

Mikaela M. Provost Department of Marine and Coastal Sciences, Rutgers University, 71 Dudley Road, New Brunswick, NJ 08901. E-mail: [email protected]

Olaf P. Jensen Department of Marine and Coastal Sciences, Rutgers University, New Brunswick, NJ

536 Fisheries | Vol. 40 • No. 11 • November 2015 The Impacts of Fishing on Hermaphroditic Species and Treatment of

Sex Change in Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 Stock Assessments

Fisheries | www.fisheries.org 537 Sex-changing fish species present unique challenges for stock assessment and management. Fishing is known to cause size at sex change to decrease and sex ratios to increasingly skew because of sex-selective fishing patterns. We show through a systematic literature review that the effects of fishing on hermaphrodite species can vary widely. Intense fishing has had no detectable effect on sex ratio and size at sex change in some stocks but a clear and dramatic impact on others. There is also substantial variation in the way stock assessments incorporate sex change. Of the 12 stock assessments of hermaphroditic fish stocks in the United States, none evaluate sex-based differences in selectivity, 10 report current sex ratio and estimate size at sex change, and only one tracks changes in both of these critical population characteristics. Despite these challenges, a global comparison of stock status suggests that the status of hermaphroditic stocks is not substantially different from that of gonochoristic species.

Impactos de la pesca en especies hermafroditas y el tratamiento del cambio de sexo en la evaluación de los stocks Las especies de peces que cambian de sexo, representan retos únicos en cuanto a la evaluación y el manejo de los stocks pesqueros. Se sabe que la pesca reduce la talla a la que los peces cambian de sexo y sesga la proporción de sexos cuando los patrones de pesca son selectivos a esta propiedad. Mediante una revisión bibliográfica se muestra que los efectos de la pesca en especies hermafroditas pueden variar ampliamente. En algunas especies, la pesca intensa ha mostrado no causar efectos detectables en la proporción de sexos ni en la edad de cambio de sexo, pero en otras los efectos son claros y drásticos. También existe una considerable variación en la forma en la que los métodos de evaluación de stocks incorporan el cambio de sexo. De las 12 evaluaciones de stocks de peces hermafroditas en los Estados Unidos de Norteamérica, ninguno consideró diferencias en la selectividad en función del sexo, 10 reportan proporción de sexos y estiman la talla a la cual ocurre el cambio de sexo y sólo en uno se rastrean los cambios en estas dos características poblacionales clave. A pesar de estos retos, una comparación global de los stocks de peces hermafroditas sugiere que su estado de explotación no es distinto que el de las especies gonocóricas.

Les impacts de la pêche sur les espèces hermaphrodites et le traitement du changement de sexe dans l’évaluation des stocks Les espèces de poissons hermaphrodites représentent des défis uniques pour l’évaluation et la gestion des stocks. La pêche est connue pour causer une diminution des changements de sexe selon la taille et fausser de plus en plus le ratio des sexes à cause des modes de pêche sélectifs en fonction du sexe. Nous montrons à travers une revue systématique de la littérature que les effets de la pêche sur les espèces hermaphrodites peuvent varier considérablement. La pêche intensive n’a eu aucun effet détectable sur le ratio des sexes et le changement de sexe selon la taille dans certains stocks, mais un impact évident et dramatique sur d’autres. Il existe aussi des variations importantes dans la façon dont les évaluations des stocks prennent en compte le changement de sexe. Sur les douze évaluations des stocks de poissons hermaphrodites aux États-Unis, aucune d’elles n’évalue les différences fondées sur le sexe en matière de sélectivité, dix font état d’un ratio des sexes et évaluent le sexe selon la taille et une seule analyse les changements de ces deux caractéristiques principales de cette population critique. Malgré ces défis, une comparaison globale de l’état des stocks suggère que l’état des stocks hermaphrodites ne diffère pas sensiblement de celle des espèces gonochoriques.

INTRODUCTION structure in two specific ways: fishing can cause a greater skew in sex ratios and/or a decrease in the size or age at sex Sex change is a life history strategy found in a wide variety change. Then, we summarize how sex change is addressed in of fish species, including many species of commercial and stock assessments and briefly review the variation in methods recreational importance. Numerous studies have examined the for calculating biological reference points in hermaphroditic implications of sex change for fitness (Ghiselin 1969; Warner stocks in the United States. Contrary to concerns in the literature

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 et al. 1975), but the implications of sex change for stock that hermaphroditic fish are particularly vulnerable to fishing, assessment and fishery management are less clearly understood. we show that the stock status of hermaphroditic species is not Previous reviews of hermaphroditism in fishes have focused on substantially different from the stock status of gonochoristic the theoretical advantages of changing sex from female to male (non-sex-changing) species and that the effects of fishing on sex (protogyny) and from male to female (protandry; Ghiselin 1969; ratio and size or age at sex change cannot be generalized to all Ross 1990) and the prevalence of these unique reproductive hermaphroditic fish species. strategies in different ecological contexts throughout extinct and extant fishes (Smith 1975). Recent studies suggest that the VULNERABILITY OF HERMAPHRODITIC timing of sex change is very flexible; some hermaphroditic FISHES TO FISHING species are capable of changing sex earlier or later in response to local fluctuations in the population sex and size ratio, and Differences in life history between hermaphroditic and select species may even change sex multiple times (Munday et gonochoristic fish species suggest that hermaphroditic species al. 2006). are especially vulnerable to fishing (Alonzo et al. 2008). In stock This review of hermaphroditism focuses on the effect that assessments of gonochoristic fish species, sex ratio is often not fishing has had on sex-changing fish species and highlights evaluated and spawning stock biomass is calculated from mature the variation in how stock assessments currently address sex female biomass alone. This simplification is reasonable if the change. First, we review why hermaphroditic fish species fishery impacts both sexes equally or if male abundance does may be especially vulnerable to fishing. Second, we show that not limit fertilization or reproductive output. In sex-changing fishing can impact hermaphroditic life history and population species, however, there is reason to suspect that fishing mortality does not impact males and females equally. Size selectivity,

538 Fisheries | Vol. 40 • No. 11 • November 2015 either inherent to the fishing gear or a result of management fishing vulnerability for two reasons. First, faster-growing fish actions such as size limits, translates into sex-based differences grow into vulnerable size ranges more quickly than slow- in fishing mortality rate for fish species where the sex ratio growing individuals; therefore, they are more susceptible changes with size. Change in sex ratio with size is a common to being captured (Sinclair et al. 2002). Individuals of feature of hermaphroditic fishes and also occurs in gonochoristic hermaphroditic species typically experience a burst in growth species with sexually dimorphic growth; for example, Summer rate immediately after sex change; for example, parrotfishes Flounder Paralichthys dentatus, (Morson et al. 2012). Size (family Scaridae; Munday et al. 2004) and Circle-Cheek Wrasse or age selectivity is often accounted for in stock assessments, Halichoeres miniatus (Munday et al. 2009). It is hypothesized but selectivity that varies by sex is rarely addressed. Fishing that this burst in growth increases their ability to compete patterns that disproportionately target one sex over the other can with existing males. Faster growth rate may push transitioning potentially have serious consequences for population growth, individuals into size classes vulnerable to fishing (such as a legal fertilization, and stability over time (Alonzo et al. 2008) and are catch size) more quickly than non-sex-changing individuals. important to consider in stock assessments of hermaphroditic Second, faster-growing fish have greater energy requirements fish species. In addition to sex ratios that vary by size, many and must forage more aggressively than slow-growing fish and, other characteristics of hermaphroditic fish species suggest that therefore, are expected to bite baited hooks more frequently sex-selective fishing is a common occurrence in fisheries of (Sutter et al. 2012). hermaphroditic species. Although the exact mechanism is unknown, there is Differences in sex-specific behavior often cause one sex to evidence that hermaphroditic species that aggregate during be more vulnerable to fishing. Aggressive behavior is known spawning tend to have increasingly skewed sex ratios after to increase catchability in gonochoristic fish species (Sutter et prolonged periods of fishing (Carter et al. 1994; de Mitcheson et al. 2012) and will likely have a similar effect in hermaphroditic al. 1994; Coleman et al. 1996). Gag M. microlepis and Scamp, species. In many protogynous species, male fish exert aggressive both protogynous hermaphrodites that form large spawning behavior while defending territories (Warner et al. 1975; aggregations, demonstrated a precipitous drop in the proportion Hoffman 1985) or guarding female mates (Frickle and Frickle 1977; Moyer and Nakazono 1978). Observations by Gilmore and Jones (1992) found that in Scamp Mycteroperca phenax, Differences in sex-specific behavior a protogynous hermaphrodite in the Gulf of Mexico, males often cause one sex to be more had greater overall movement, displayed more aggressive behavior (e.g., lunging, attacking), and (anecdotally) had a vulnerable to fishing. higher propensity to bite hooks compared to females of a similar length. The authors hypothesized that male Scamp have higher of males (from 17% to 1% and 36% to 18%, respectively) fishing mortality because of their aggressive behavior as seen over a span of 25 years of exploitation in the Gulf of Mexico in underwater videos. Aggression levels also differ between (Coleman et al. 1996). In contrast, Red Grouper Epinephelus individuals based on social status within social aggregations; morio, a nonaggregating protogynous hermaphrodite with an in protogynous hermaphrodites, dominant males express the overlapping range, showed little change in sex ratio over the most aggression (Layton and Fulton 2014). There are, however, same time period and under similar fishing conditions. There are exceptions such as Red Porgy Pagrus pagrus, a protogynous many reasons that may explain this phenomenon; for example, hermaphrodite located in the Gulf of Mexico and South Atlantic. behavior could be sex-specific in spawning aggregations (e.g., In this species, males show little aggression, and male and Taborsky 1994), lending one sex to be more vulnerable to female fish appear to be captured proportionally, suggesting capture, or spawning aggregations could be predominately that males and females are equally vulnerable to hook-and-line composed of a single sex (de Mitcheson et al. 1994) so that only fishing gear (DeVries 2007). the aggregating sex is exposed to intense fishing. Hermaphroditic fishes may also be vulnerable to fishing Changes in cohort strength may also play an important role because fertilization is dependent upon complex social in sex ratio fluctuations over time. If fishing disproportionately Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 structures, such as harems, formed during spawning. Social removes the terminal sex—that is, males in protogynous fish structures help facilitate mate choice during the spawning season species and females in protandrous species—leading to an (Moyer and Nakazono 1978; Nemtzov 1985; McCormick et increasingly skewed sex ratio, then a large cohort entering the al. 2010). Frequent removal of large, dominant individuals by population as the initial sex will push the skewed sex ratio even fishing may interrupt these social hierarchies. It is possible that further. Variability of cohort size could translate to variability in such removal could delay female reproduction and negatively recruitment because highly skewed sex ratios may lead to lower impact fertilization rates, especially in species with a short fertilization rates. spawning season and narrow window of opportunity for mating. In Red Hind Epinephelus guttatus, for example, small Effect of Skewed Sex Ratios and Decreases in Size or Age groups consisting of a single male and multiple females come at Sex Change on Fertilization Rates together during a two-week spawning period (de Mitcheson Increasingly skewed sex ratios will, at some point, result in et al. 1994). Exposure to intense fishing pressure during this reduced fertilization rates and, consequently, slow population narrow timeframe could result in rapid removal of large, male growth. Though the impact of incremental changes in the individuals. In situations similar to this, females may not have sex ratio on fertilization rates is not well understood in fish, sufficient time to form new spawning groups or, alternatively, there are several examples of reduced fertilization rates in reverse their sex and take advantage of mating opportunities marine invertebrates. A heavily male-biased fishery for blue with the recently abandoned females. crab Callinectes sapidus in the Chesapeake Bay is thought Sexually dimorphic growth, a characteristic of some to have resulted in decreased fertilization rates because of gonochoristic and many hermaphroditic fishes, likely increases sperm limitation (Hines et al. 2003). Similarly, in a population

Fisheries | www.fisheries.org 539 of Caribbean sea urchin Diadema antillaru, sex ratios were species—leaving behind a greater proportion of female fish. This dramatically skewed after a severe mortality event (Lessios phenomenon has been documented in Steentjie Spondyliosoma 1988). Although female urchin body size and egg production per emarginatum (Tunley et al. 2009), Red Hind (Beets and female greatly increased because of low population density after Friedlander 1998), and Snowy Grouper Epinephelus niveatus the die-off, the number of fertilized zygotes produced per female (Wyanski et al. 2000; Table 1). did not increase because female fertilization was drastically reduced in the absence of sufficient numbers of male sea urchins Literature Review (Levitan 1992). Severe reductions in the proportion of male fish Based on a review of published literature, we determined in populations of protogynous hermaphrodites may have similar the degree of support for each hypothesis regarding the effects effects (Alonzo and Mangel 2004), and the impact of sex ratio of fishing on hermaphroditic fish species. A systematic literature on fertilization rate remains one of the critical uncertainties in search was carried out using Web of Science (thomsonreuters. understanding the population dynamics of hermaphroditic fishes com/thomson-reuters-web-of-science) for publications published (Brooks et al. 2008). between 1980 and 2014. There was no restriction of origin One potential side effect of male scarcity in protogynous and language. Search terms included all combinations of species is reduced genetic diversity (Chapman et al. 1999) and “fish” and (“sex change” or “sex ratio”) and (“protogyny” or possible Allee effects: the phenomenon of negative population “hermaphroditism” or “hermaphrodite”). Publications found growth at low population size (Allee 1931). Even if fertilization in Web of Science search results were screened and restricted rates remain high during times of male scarcity, the bottleneck to studies that measured size or age at sex change and/or the of genetic material through a relatively small number of males sex ratio in populations exposed to different levels of fishing. will decrease the population’s overall genetic diversity and Some studies assessed differences among populations that may reduce the population’s ability to adapt to environmental were exposed to different levels of fishing; for example, variability and novel pathogens. These less visible genetic comparisons between populations inside and outside of a bottleneck effects can occur even while fertilization rates remain marine protected area (Götz et al. 2008). Other publications high (Levitan 1992). may have tracked changes in size or age at sex change and/ or the sex ratio within a single population that was exposed to HOW DOES FISHING AFFECT HERMAPHRODITIC increasing (or decreasing) levels of fishing over a long period FISH SPECIES? WHAT HAVE WE LEARNED of time (e.g., Hamilton et al. 2007). Studies that carried out in FROM DECADES OF FIELD OBSERVATIONS? situ removal experiments to simulate the process of fishing were Fishing is likely to impact hermaphroditic fish species in one of two ways: (1) size or age at sex change will Table 1. Summary of results from the systematic literature review showing the number of publications and species in support of hypothesis 1, size decrease with greater fishing intensity, or (2) sex ratios or age at sex change decreases under heavy fishing conditions (H1); will become increasingly skewed. hypothesis 2, sex ratios increasingly skew under heavy fishing conditions (H2); both hypotheses 1 and 2 (H1 and H2); and those where fishing had no Hypothesis #1: Heavy Fishing Causes a Downward effect on size or age at sex change and sex ratios (NC). Shift in Age or Size at Sex Change Number of Number of Alternatively, the sex ratio may change very little or Hypothesis publications species References even remain constant under intense fishing conditions in support examined because of decreases in size or age at sex change. In H1 (size at 5 5 Mariani et al. (2013), Mackie sex change (2003), some protogynous species, females will compensate for decreases) Platten et al. (2002), Warner the loss of male individuals by changing sex at earlier and Swearer (1991), Götz et al. ages and smaller sizes. Sex change is often a socially (2008) mediated process (Warner and Swearer 1991; Benton H2 (sex 7 10 Coleman et al. (1996), Tunley et and Berlinsky 2006), and the process of sexual transition ratio al. (2009), Beets and Fried- acts as a compensatory mechanism to maintain an changes) lander (1998), Hawkins and Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 Roberts (2003), McGovern optimal sex ratio (Shapiro 1979). This phenomenon has et al. (1998), Wyanski et al. been observed in laboratory experiments (Benton and (2000), Buxton (1993) Berlinsky 2006) and in multiple field studies (Table 1). Decreases in the size or age at sex change can occur over periods as long as two to three decades; for example, H1 and 5 5 S. A. Shepherd et al. (2010), H2 (both Hamilton et al. (2007), Mariani in California Sheephead Semicossyphus pulcher, a effects et al. (2013), Buxton (1993), protogynous hermaphrodite in south-central California, happen) Harris and McGovern (1997) female size at sex change decreased between 72 and 168 mm over approximately 25 years (Hamilton et al. 2007). Decreases can also occur in as short as a few weeks; for example, in a population of protogynous Blue-Headed Wrasse Thalassoma bifasciatum in the U.S. Virgin Islands, size at sex change began decreasing within two weeks of heavy exploitation (Warner and Swearer 1991). NC (no 2 6 Hawkins and Roberts (2003), change in Adams et al. (2000) Hypothesis #2: Heavy Fishing Results in sex ratio Increasingly Skewed Sex Ratios and size Sex-selective fishing patterns disproportionately at sex target the larger sex—that is, males in protogynous change)

540 Fisheries | Vol. 40 • No. 11 • November 2015 also included in this literature review. Reference lists in each considerably less time reproducing as females if male fishing publication found through Web of Science and included here mortality is disproportionately high because females may were searched to find additional relevant studies. In total, 244 undergo sex reversal earlier than usual. Without accounting for publications were screened from Web of Science search results, the accelerated loss of female biomass to sex change, population and 735 publications were screened from bibliographies of dynamics models may overestimate spawning stock biomass qualifying papers found through Web of Science. Sixteen papers (Alonzo et al. 2008). Second, the potential for lower fertilization that studied 26 hermaphroditic species presented quantitative rates during times of male scarcity complicates recruitment estimates of changes in sex ratio or size or age at sex change. calculations in population dynamics models (Brooks et al. Publications found through the systematic literature review 2008). If the potential for sperm limitation is unknown, which measured fishing pressure in a variety of ways. For example, is likely true for most managed hermaphroditic stocks, then

Hamilton et al. (2007) verified that there were changes in estimates of biological reference points (e.g., BMSY) are better exploitation level by comparing estimates of annual survivorship predicted when recruitment is measured as a function of both between historical and recent samples, and Tunley et al. (2009) male and female spawning stock biomass (Brooks et al. 2008) were able to approximate fishing intensity by landings from rather than just female biomass. Ultimately, however, the effects recreational boats and shore anglers. For the purpose of this of fishing on hermaphroditic population dynamics depends on literature review, the type of fishing pressure reported in each the disproportionate fishing mortality rate of males and how study is not relevant, only that size or age at sex change and/or strong of an effect male removal has on female sex change and sex ratios were assessed at different levels of fishing. The phrase fertilization rates (Alonzo and Mangel 2005). “fishing intensity” is used throughout this article to reference We reviewed the treatment of sex change in stock the general fishing level and covers such terms as exploitation, assessments for nine hermaphrodite fish species consisting of harvest, and fishing mortality. 12 different stocks in the Mid-Atlantic, South Atlantic, and Gulf Each eligible study was identified as showing support for of Mexico and one species from California. Species included either one of the two aforementioned hypotheses or for neither Black Grouper M. bonaci, Black Sea Bass Centropristis hypothesis: size at transition decreases under high fishing striata, Gag, Hogfish Lachnolaimus maximus, Red Grouper, intensity (H1) or sex ratios shift—that is, in favor of females in Red Porgy, Snowy Grouper, Yellowedge Grouper Epinephelus protogynous species and males in protandrous species—while flavolimbatus, and California Sheephead (Table 2). In all of under intense fishing pressure (H2; Table 1). For some species, these fisheries, fish are primarily captured by hook-and-line gear fishing led to changes in both the sex ratio and size or age at or traps. Overall, we found that there is considerable variation in sex change (H1 and H2). Lastly, some studies found that fishing the data reported that is needed to assess the impact fishing has had no effect on the sex ratio and size or age at sex change (no on size or age at sex change and sex ratio. change, NC). The majority of stock assessments did not track changes We found more studies documenting shifts in the sex ratio in sex ratio or size at sex change, and none reported estimates under intense fishing (H2, 7/16) and slightly fewer studies of sex selectivity in the fishery (Table 2). Of the 12 stock showing fishing caused decreases in size or age at sex change assessments we reviewed, 10 included both male and female (H1, 5/16; Table 1). The seven publications in support of biomass in spawning stock biomass calculations, as suggested H2 reported that sex ratios were increasingly skewed in 10 by Brooks et al. (2008) for more reliable spawning biomass hermaphroditic species. Five publications found in support estimates in hermaphroditic species. Most assessments, 10 of 12, of H1 showed that size or age at sex change decreased in five also reported the current sex ratio and estimates of size or age hermaphroditic species. There were five publications that at sex change; however, only 4 of these 10 assessments reported showed both sex ratios and size or age at sex change shifted whether the size or age at sex change has changed over time, with increasing fishing intensity in the same species (H1 and and only 2 of the 10 assessments evaluated changes in the sex H2, 5/16). Two of the 16 publications specifically documented ratio over time. that fishing pressure did not affect sex ratio or size or age at sex We found there to be some variation in the way reference change in six species (NC). points were calculated among stock assessments, especially Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 It should be noted that not every study found through the between stocks from different management regions (Table literature review assessed both sex ratio and size or age at sex 3). Biological reference points are a critical part of stock change. A lack of support for one hypothesis by one study for a assessment because they are used to guide fishery management certain species should not be interpreted as evidence against that objectives—that is, how hard to fish—and are calculated based hypothesis. For example, Coleman et al. (1996) tracked changes on biological characteristics of the species (Quinn and Deriso in sex ratio over time in three groupers in the Gulf of Mexico. 1999). Incorporating a sex change parameter into reference The authors found that sex ratio changed for two of the three point calculations is expected to lead to more reliable estimates species but reported no information on changes in size or age at of spawning stock biomass (Alonzo et al. 2008). Most stock sex change for all three species; therefore, we cannot make any assessments from the South Atlantic, specifically Red Grouper, conclusions regarding the support for H2 for each species. Even Snowy Grouper, Red Porgy, and Gag, estimated reference though each study did not fully address both sex ratio and size points based on maximum sustainable yield (MSY) from the or age at sex change, the results in Table 1 show that fishing has Beverton-Holt stock recruitment model. Transition from female not had a uniform effect on hermaphroditic fish species. to male was modeled using logistic regression, which was input to the statistical catch-at-age model as proportion female at age. HOW DO STOCK ASSESSMENTS Spawning stock biomass was then computed using both female ACCOUNT FOR SEX CHANGE? and male biomass. In the Gulf of Mexico, there was more The effect of skewed sex ratios and decreases in the size or variation among assessments: reference points for both Red age at sex change on fertilization rates pose specific challenges Grouper and Yellowedge Grouper were also based on MSY from for stock assessment. First, protogynous fishes could spend a Beverton-Holt stock recruitment model. Reference points for

Fisheries | www.fisheries.org 541 Table 2. Treatment of sex change in stock assessments of protogynous hermaphrodite fish species in the United States. Measures Spawning Reports Change in Reports sex or esti- stock change in the age (or Family Stock and stock locations ratio at age mates of Reference biomass proportion size) at sex (or size) sex selec- (SSB) male change tivity Serranidae Black Sea Bass Male + Yes Not Not Not SAFMC (2013b) Centropristis striata female reported reported reported South Atlantic Black Sea Bass Male + Yes Not Not Not G. R. Shepherd and Centropristis striata female reported reported reported Nieland (2012) Mid-Atlantic Black Grouper Male + Yes Not Not Not SAFMC (2010b) Mycteroperca bonaci female reported reported reported South Atlantic, Gulf of Mexico Gag Grouper Female Not Not Yes (no Not GMFMC (2014) Mycteroperca microlepis reported reported decrease reported Gulf of Mexico reported) Gag Grouper Male + Yes Not Yes (no Not SAFMC (2006) Mycteroperca microlepis female reported decrease reported South Atlantic reported) Red Grouper Male + Yes Yes Yes (no Not SAFMC (2010a) Epinephelus morio female decrease reported South Atlantic reported) Red Grouper Female Yes Not Yes (no Not GMFMC (2009) Epinephelus morio reported decrease reported Gulf of Mexico reported) Snowy Grouper Male + Yes Not Not Not SAFMC (2013a) Epinephelus niveatus female reported reported reported South Atlantic Yellowedge Grouper Male + Yes Not Not Not GMFMC (2011) Epinephelus flavolimbatus female reported reported reported Gulf of Mexico Labridae California Sheephead Male + Not Not Not Not Alonzo et al. (2004) Semicossphus pulcher female reported reported reported reported California Hogfish Male + Yes Not Not Not SAFMC (2004) Lachnoaimus maximus female reported reported reported S. Atlantic, Gulf of Mexico Sparidae Red Porgy Male + Yes Yes Not Not SAFMC (2012) Pagrus pagrus female reported reported South Atlantic

Gag were based on yield per recruit, and for Black Grouper they One reason why hermaphroditic stocks may not be worse

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 were based on spawning potential ratio. off than gonochoristic stocks could be that some sex-changing Although we expect hermaphroditic fish species to be species compensate for sex-selective fishing. Hermaphroditic especially vulnerable to fishing, as discussed previously, fishes that maintain an optimal sex ratio are expected to be less hermaphrodite gagstock status, in general, is no better or worse vulnerable to fishing based on simulation studies (Huntsman than stock status of gonochoristic stocks (Figure 1). By plotting and Schaaf 1994). This suggests that when sex change is biological reference points from 189 fish stocks in the RAM affected by exogenous social cues, such as the local sex ratio Legacy Stock Assessment Database (Ricard et al. 2012), we (Munday 2002) or density (Lutnesky 1994), the species may compared biomass relative to the level of biomass at MSY be more resilient to fishing pressure compared to species

(B:BMSY) and exploitation rate relative to the exploitation rate at where sex change is triggered through endogenous cues; for

MSY (U:UMSY) for gonochoristic (n = 178) and hermaphroditic example, size or age of fish (Alonzo and Mangel 2005). In the (n = 11) stocks. California Sheephead were not included in preceding literature review, it appears that some species are this comparison because reference points have not yet been able to compensate for sex-selective fishing (H1), whereas other estimated. The biomass of 8 out of the 11 hermaphroditic stocks species are not able to maintain sex ratios (H2). Hawkins and

(73%) was below BMSY compared to 57% for gonochoristic Roberts (2003) found that high fishing intensity did not always species, a difference that is not statistically significant (χ2 = lead to skewed sex ratios and smaller size at sex change among 0.53, df = 1, P = 0.47). The current exploitation rate was above different parrotfishes species; similarly, Coleman et al. (1996)

UMSY for 5 of the 11 hermaphroditic stocks (45%) compared showed that fishing lead to highly skewed sex ratios in some to 50% for gonochoristic species, a difference that is also not Serranidae species but not others. Maintaining optimal sex ratios statistically significant (χ2 = 0.01, df = 1, P = 0.97). presumably helps to keep fertilization rates high (Huntsman

542 Fisheries | Vol. 40 • No. 11 • November 2015 Table 3. Summary of the types of stock assessment models, methods for calculating biological reference points, and methods of incorporating sex change into stock assessment for hermaphroditic stocks in the United States. MSY is maximum sustainable yield, and YPR is yield per recruit. Stock and management Method for estimating Method of incorporating sex change Assessment model type region reference points (if any) Black Sea Bass, Beaufort assessment Benchmarks were calculated Sex ratio is estimated using South Atlantic, model, a statistical catch- based on MSY estimates from logistic regression; spawning stock SAFMC (2013b) at-age model the Beverton-Holt stock-re- biomass is computed using female cruitment curve. mature biomass. Black Sea Bass, Statistical length-based YPR Spawning stock biomass is computed mid-Atlantic, model (SCALE) using both male and female biomass. G. R. Shepherd and Nieland (2012) Black Grouper, Statistical catch-at-age Benchmarks are based on Sex ratio is estimated using logistic South Atlantic, model (ASAP2) spawning potential ratio. regression; spawning stock biomass Gulf of Mexico, is computed using female and male AFMC (2010b) mature biomass.

Gag, Stock synthesis assessment Benchmarks are based on YPR NA Gulf of Mexico, model estimated from stock synthesis GMFMC (2014) (for females only).

Gag, Beaufort assessment Benchmarks were calculated Sex ratio is estimated using logistic South Atlantic, model, a statistical catch- based on MSY estimates from regression; spawning stock biomass SAFMC (2006) at-age model the Beverton-Holt stock-re- is computed using female and male cruitment curve. mature biomass.

Red Grouper, Beaufort assessment Benchmarks were calculated Sex ratio is estimated using logistic South Atlantic, model, a statistical catch- based on MSY estimates from regression; spawning stock biomass SAFMC (2010a) at-age model the Beverton-Holt stock-re- is computed using female and male cruitment curve. mature biomass. Red Grouper, Age-structured forward Benchmarks were calculated NA Gulf of Mexico, projection model based on MSY estimates from GMFMC (2009) the Beverton-Holt stock-recruitment model. Snowy Grouper, Beaufort assessment Benchmarks were calculated Sex ratio is estimated using logistic South Atlantic, model, a statistical catch- based on MSY estimates from regression; spawning stock biomass SAFMC (2013a) at-age model the Beverton-Holt stock-re- is computed using female and male cruitment curve. mature biomass. Yellowedge Grouper, Statistical catch-at-age Benchmarks were calculated Spawning stock biomass is computed Gulf of Mexico, based on MSY estimates from using both male and female biomass. GMFMC (2011) the Beverton-Holt stock-recruitment curve. California Sheephead, Stock synthesis length- Benchmarks were based on Developed two models: California, based model changes in spawning potential (1) Two-sex model—All individuals Alonzo et al. (2004) ratio. recruit to population as female, become mature with probability at each length-at-age, then become male with a different probability at each

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 length-at-age. 2) Single-sex model—Only tracks females. The two models differ in the maturity and fecundity functions. Hogfish, South Atlantic, REEFS, a sex-differentiated, Reference points are based A parameter representing sex ratio at SAFMC (2004) age-structured, stochastic on YPR. age is included. length-based population simulation model Red Porgy, South Atlantic, Beaufort assessment Benchmarks were calculated Sex ratio at age is estimated using SAFMC (2012) model, a statistical catch- based on MSY estimates from logistic regression; spawning stock at-age model the Beverton-Holt stock- biomass is computed using female recruitment curve. and male mature biomass.

and Schaaf 1994), and it may be that those hermaphroditic fish species is not significantly different from that of gonochoristic species able to shift size or age at sex change are better equipped species, as fishing intensity rises hermaphroditic species may to withstand moderate fishing intensity. not be able to compensate enough for sex-selective fishing, and Accurately assessing the impact of fishing will require stock assessments will need to incorporate not only size and age tracking changes in both male and female biomass over time, but also sex into stock assessment models. because male biomass may be just as important as female In conclusion, it is difficult to generalize the effects of biomass in populations of hermaphrodite species (Brooks et fishing on hermaphroditic species as seen from the literature al. 2008). Even though overall stock status of hermaphroditic review. For many hermaphroditic species, fishing has impacted

Fisheries | www.fisheries.org 543 Figure 1. Biomass and exploitation rates relative to levels expected at maximum

sustainable yield (BMSY and UMSY, respectively) for hermaphroditic (filled circles, n = 11) and gonochoristic (open circles, n = 178) fish stocks from the RAM Legacy Stock Assessment Database (Ricard et al. 2012).

the sex ratio, size or age at sex change, or both. A lack of Alonzo, S. H., and M. Mangel. 2004. The effects of size-selective consistency in fishing’s impact is likely because fishing’s fisheries on the stock dynamics of and sperm limitation in sex- changing fish. Fishery Bulletin 102:1–13. influence is highly species specific and in part depends on life ———. 2005. Sex-change rules, stock dynamics, and the performance history characteristics other than sex change itself. The variation of spawning-per-recruit measures in protogynous stocks. Fish- in fishing’s influence on sex ratio and size or age at sex change, ery Bulletin 103:229–245. Beets, J., and A. Friedlander. 1998. Evaluation of a conservation strat- in addition to the variation in how stock assessments address egy: a spawning aggregation closure for Red Hind, Epinephelus sex change, may suggest why hermaphroditic species are not guttatus, in the U.S. Virgin Islands. Environmental Biology of significantly worse off than gonochoristic species. A nuanced Fishes 55:91–98. understanding of the impacts of fishing on sex-changing fishes Benton, C. B., and D. L. Berlinsky. 2006. Induced sex change in black Sea Bass. Journal of Fish Biology 69:1491–1503. requires species-specific studies of sex change and life history, Brooks, E. N., K. W. Shertzer, T. Gedamke, and D. S. Vaughan. 2008. time series of sex ratio and size or age at sex change, and Stock assessment of protogynous fish: evaluating measures of estimation of sex selectivity of fishing gear. spawning biomass used to estimate biological reference points. Fisheries Bulletin 106:12–23. Buxton, C. D. 1993. Life history changes in exploited reef fishes on ACKNOWLEDGMENTS the east coast of South Africa. Environmental Biology of Fishes We are grateful for comments and assistance from Phil 36:47–63. Carter, J., F. J. Marrow, and V. Pryor. 1994. Aspects of the ecology Neubauer, Anthony Vastano, Kyle Shertzer, and participants in and reproduction of Nassau Grouper, Epinephelus striatus, off the workshop “Modeling Protogynous Hermaphrodite Fishes” the coast of Belize, Centeral America. Proceedings of the 43rd Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 chaired by Jessica Coakley. Gulf and Caribbean Fisheries Institute 65–111. Chapman, R. W., G. R. Sedberry, C. C. Koenig, and B. M. Eleby. 1999. Stock identification of Gag,Mycteroperca microlepis, along the FUNDING southeast coast of the United States. Marine Biotechnology This work was supported by the National Oceanic and 1:137–146. Coleman, F. C., C. C. Koenig, and L. A. Collins. 1996. Reproductive Atmospheric Administration’s Research Set Aside Program and styles of shallow-water groupers (Pisces: Serranidae) in the New Jersey Sea Grant. eastern Gulf of Mexico and the consequences of fishing spawning aggregations. Environmental Biology of Fishes 47:129–141. REFERENCES de Mitcheson, Y. S., A. Rosario, and A. Roman. 1994. Reproduction in an aggregating grouper, the Red Hind, Epinephelus guttatus. Adams, S., B. D. Mapstone, G. R. Russ, and C. R. Davies. 2000. Geo- Environmental Biology of Fishes 41:269–286. graphic variation in the sex ratio, sex specific size, and age struc- DeVries, D. A. 2007. No evidence of bias from fish behavior in the ture of Plectropomus leopardus (Serranidae) between reefs selectivity of size and sex of protogynous Red Porgy (Pagrus open and closed to fishing on the Great Barrier Reef. Canadian pagrus, Sparidae) by hook-and-line gear. Fisheries Bulletin Journal of Fisheries and Aquatic Sciences 57:1448–1458. 105:582–587. Allee, W. C. 1931. Animal aggregations, a study in general sociology. Frickle, H., and S. Frickle. 1977. Monogamy and sex change by ag- University of Chicago Press, Chicago. gressive dominance in coral reef fish. Nature 266:830–832. Alonzo, S. H., T. Ish, M. Key, A. D. MacCall, and M. Mangel. 2008. The Ghiselin, M. T. 1969. The evolution of hermaphroditism among ani- importance of incorporating protogynous sex change into stock mals. The Quarterly Review of Biology 44:189–208. assessments. Bulletin of Marine Science 83:163–179. Gilmore, R. G., and R. S. Jones. 1992. Colar variation and associated Alonzo, S. H., M. Key, T. Ish, and A. D. MacCall. 2004. Status of the behavior in the epinepheline groupers, Mycteroperca microlepis California Sheephead (Semicossyphus pulcher) stock. California (Goode and Bean) and M. phenax Jordan and Swain. Bulletin of Fish and Game 1–118. Marine Science 51:83–103.

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Fisheries | www.fisheries.org 545 FEATURE

The Role of Introduced Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 Populations in the Management and Conservation of Least Chub

546 Fisheries | Vol. 40 • No. 11 • November 2015 The Role of Introduced Populations in the Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 Management and

Paul D. Thompson Utah Division of Wildlife Resources, 515 East 5300 South, Ogden, UT 84405. E-mail: [email protected]

Conservation of P. Aaron Webber U.S. Fish and Wildlife Service, Bethel, AK

Cassie D. Mellon Least Chub U.S. Bureau of Land Management, West Valley, UT

Fisheries | www.fisheries.org 547 Native fishes continue to decline in abundance and distribution. One common practice to ensure native fish persistence has been to introduce fish into new habitats. Though these introductions reduce the risk of extinction, often the introduced populations are not considered in the Endangered Species Act (ESA) listing status in the same manner as extant populations. One instance where they were considered was when Least Chub Iotichthys phlegethontis were introduced into 23 locations within the Bonneville Basin between 2005 and 2013. In 2014, 10 of these populations were considered successful and were evaluated with the six remaining extant populations by the U.S. Fish and Wildlife Service. In this instance, Least Chub were removed from the candidate list in part due to these introduced populations, which increased the resiliency, redundancy, and representation for this species to persist now and into the foreseeable future. The creation of introduced fish populations is a conservation practice that needs to be considered as a potential tool for fisheries managers, not only to ensure persistence, but also to preclude the need for federal listing under ESA.

El papel de poblaciones introducidas en el manejo y conservación del Least Chub La abundancia y distribución de los peces nativos continúan declinando. Una práctica común para asegurar la persistencia de los peces nativos ha sido introducirlos en nuevos hábitats. Si bien estas introducciones reducen el riesgo de extinción, a veces las poblaciones introducidas, en comparación a las poblaciones ya existentes, no son consideradas de la misma forma dentro de los listados del AEA (Acta de Especies Amenazadas). La única instancia en la que fueron consideradas así fue cuando el Least Chub (Iotichthys phlegethontis) fue introducido en 23 sitios dentro de la cuenca Bonneville entre 2005 y 2013. En 2014, 10 de estas poblaciones fueron consideradas como un éxito y el Departamento de Pesca y Vida Salvaje (DPVS) de los EE.UU. se encargó de evaluarlas junto con las seis poblaciones restantes que existían. En este caso, el Least Chub fue removido de la lista de especies amenazadas en parte debido a dichas poblaciones introducidas, las cuales incrementaron la resiliencia, redundancia y representatividad de la especie en pos de su persistencia presente y futura. La creación de poblaciones introducidas de peces es una práctica de conservación que debe ser considerada como una potencial herramienta para los manejadores de pesquerías, no solo con el fin de asegurar la persistencia de las especies sino también como medida preventiva para evitar la necesidad de listarlas en los elencos del AEA.

Le rôle des populations introduites dans la gestion et la conservation des Iotichthys Phlegethontis (Least Chub) L’abondance et la distribution de poissons indigènes continuent à décliner. Une pratique courante pour assurer la persistance de poissons indigènes a été d’introduire les poissons dans de nouveaux habitats. Bien que ces introductions réduisent le risque d’extinction, bien souvent les populations introduites ne sont pas prises en compte dans la liste des espèces en danger de l’ESA de la même manière que les populations existantes. l’iotichthys phlegethontis (Least Chub) a été introduit dans 23 endroits dans le bassin de Bonneville entre 2005 et 2013 et est l’un des cas où cela a été pris en considération. En 2014, 10 de ces populations ont bien réussi et ont été évaluées avec les six autres populations existantes par le Fish and Wildlife Service des États-Unis (USFWS). Dans ce cas, les Iotichthys phlegethontis (Least Chub) ont été retirés de la liste des candidats, en partie parce que ces populations introduites ont augmenté leur résilience, leur redondance et leur représentation et permis à cette espèce de persister maintenant et dans un avenir prévisible. La création de populations de poissons introduites est une pratique de conservation qui doit être considérée comme un outil potentiel pour les gestionnaires des pêches, non seulement pour assurer la persistance, mais aussi pour empêcher la nécessité de tenir une liste fédérale en vertu de l’ESA.

INTRODUCTION Least Chub Iotichthys phlegethontis populations have declined as a result of these same factors. The Least Chub is Native fishes have declined steadily in abundance and Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 a minnow endemic to the Bonneville Basin of Utah, and they distribution across western North America in the 20th are the only species of the genus Iotichthys. Least Chub are century (Williams et al. 1989; Moyle and Leidy 1992), and the smallest cyprinids in Utah, reaching a maximum size of approximately one-half of the fish species listed as threatened only 76 mm total length (TL; Sigler and Sigler 1996; Figure or endangered, or being considered for listing by the U.S. Fish 1). The Least Chub are considered a short-lived species, and and Wildlife Service (USFWS), occur west of the Continental some individuals have been aged up to six years (Mills et Divide (Minckley and Douglas 1991). These declines have been al. 2004a). Least Chub mature at age 1 (Sigler and Sigler especially pronounced with species occupying desert spring 1996) and spawn intermittently throughout the late spring and habitats (Minckley and Douglas 1991). Factors attributed to the summer (Crawford 1979). Least Chub feed primarily on algae decline of desert fishes include the alteration of environmental and small invertebrates (Sigler and Sigler 1996) and persist conditions associated with water development, improper in fish communities with low species diversity. The rarity and grazing management, nonnative fish, and changes in land use imperilment of Least Chub has been recognized for more than and management, which has resulted in habitat degradation. 40 years (Table 1), and although once common in springs, lakes, Introduced nonnative fishes became established as they were streams, marshes, rivers, and ponds within the Bonneville Basin better adapted to these conditions or could outcompete the (Sigler and Sigler 1987; Page and Burr 1991), only six known native fish population. These introduced fishes contributed to the extant Least Chub populations remain (Figure 2; Bailey et al. decline of desert fish through competition, predation, and other 2005; Mock and Bjerregaard 2007). One of the most significant detrimental interactions (Meffe 1985; Minckley and Douglas factors attributed to their decline has been the introduction of 1991; Lydeard and Belk 1993; Stevens and Meretsky 2008). Western Mosquitofish Gambusia affinis. Western Mosquitofish

548 Fisheries | Vol. 40 • No. 11 • November 2015 Figure 1. Typical adult Least Chub with a female (left) and a male in spawning colors (right). Photo credit: Paul Thompson.

Table 1. Least Chub listing status and significant decisions/actions taken in relation to ESA listing. Year Agency/groupa Decision/action Reference 1972 AFS Classified as a threatened species Miller (1972) 1979 AFS Classified as a threatened species Deacon et al. (1979) 1980 USFWS Status review, insufficient data to be classified as threatened or endangered Bailey et al. (2005) species 1982 USFWS Classified as category 2 candidate species USFWS (1982) 1989 USFWS Classified as category 1 candidate species USFWS (1989) 1995 USFWS Proposed to list as endangered with critical habitat under the Endangered USFWS (1995) Species Act (ESA) 1995 Congress Listing moratorium suspended listing action Bailey et al. (2005) 1998 Signatories Developed Least Chub conservation agreement and strategy Perkins et al. (1998) of agreement 1998 Signatories Formed Least Chub Conservation Team (LCCT) Perkins et al. (1998) of agreement 2001 UDWR Considered Least Chub a conservation agreement species UDWR Sensitive Species Listb

2005 LCCT Revised 1998 Least Chub conservation strategy Bailey et al. (2005) 2007 Petitioners Petitioned Least Chub to be listed as threatened under ESA and critical habitat

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 be designated 2008 USFWS 90-Day finding concluded that petition presented substantial information in USFWS (2008) support of listing 2010 USFWS 12-Month finding identified Least Chub as a species for which listing as USFWS (2010a) endangered or threatened was warranted, but was precluded 2010 USFWS Classified as candidate species USFWS (2010b) 2011 USFWS Candidate notice of review (CNOR) maintained the species as a candidate USFWS (2011) 2012 USFWS CNOR maintained the species as a candidate USFWS (2012) 2013 USFWS CNOR maintained the species as a candidate USFWS (2013) 2014 LCCT Amended 2005 Least Chub conservation strategy LCCT (2014) 2014 USFWS Removed from list of candidates under ESA USFWS (2014b)

aAFS = American Fisheries Society; USFWS = United States Fish and Wildlife Service; Congress = United States Congress; Signatories of agreement = signatories to the Least Chub conservation agreement and strategy as of 2014 include Utah Department of Natural Resources, Division of Wildlife Resources; Bureau of Land Management; United States Fish and Wildlife Service; Bureau of Reclamation; Utah Reclamation Mitigation and Conservation Commission; Central Utah Water Conservancy District; Confederated Tribes of the Goshute Reservation; and Southern Nevada Water Authority; UDWR = Utah Division of Wildlife Resources; LCCT = Least Chub Conservation Team; Petitioners = Center for Biological Diversity, Confederated Tribes of the Goshute Reservation, Great Basin Chapter of Trout Unlimited, and Utah Chapter of the Sierra Club. bThe UDWR also considered the Least Chub a wildlife species of concern and a federal candidate species in past sensitive species lists.

Fisheries | www.fisheries.org 549 refers to both the translocation of wild fish and the stocking of hatchery Least Chub to establish new populations within their historic range. The goal of introducing additional Least Chub populations was to maintain the genetic diversity and integrity of the wild populations while reducing the extinction risk of Least Chub due to environmental stochasticity or random catastrophes (Lande 1993). Echelle (1991) recommended that 200 or more fish be utilized in a founding stock for intermittent breeders because some losses of rare alleles can occur when smaller founding stocks of less than 100 are utilized. Nuclear (amplified fragment length polymorphism) and mitochondrial genetic markers from two previous Least Chub introductions, Lucin Pond and Walter Springs, used similar donor numbers of n = 131 and n = 230 fish, respectively. Mock and Miller (2005) found that these populations were similar to their donor populations with respect to genetic diversity measures and there was no evidence of reduced diversity or severe bottlenecks in these populations. The introduction of Least Chub into additional locations began as early as 1979, with nine introductions documented from 1979 to 2004. These early introductions were primarily opportunistic and though some persisted for a couple years, they were largely unsuccessful. Since 2005, 23 Least Chub introductions have been attempted and 12 reproducing and recruiting Least Chub populations have been established. The objectives for introducing Least Chub populations were to (1) increase redundancy and resiliency and prevent possible Figure 2. Location of extant and successful introduced Least Chub extinction of the remaining six extant populations and (2) populations within three geographic management units in the Bon- preclude the need for federal listing under the Endangered neville Basin, Utah. Species Act (ESA). Objectives specific to this article were to demonstrate how introduced populations can be a conservation are known to be aggressive and predate on eggs and young of tool for fisheries managers and review how the USFWS viewed other fishes (Meffe 1985; Sigler and Sigler 1987; Mills et all. introduced populations in the Least Chub listing decision under 2004b). ESA. In 1998, a Least Chub Conservation Agreement and METHODS Strategy (LCCAS) was developed that outlined specific actions and strategies needed to ensure the persistence of the species Study Site (Perkins et al. 1998). The Least Chub Conservation Team Introduced populations of Least Chub were required to be (LCCT) was formed and revised the LCCAS in 2005 (Bailey within the historic range of the species, the Bonneville Basin. et al. 2005) and in 2014 (LCCT 2014; Table 1). The genetic Many larger water bodies were excluded due to the prior characterization of the six extant wild Least Chub populations establishment of species that are known to be noncompatible indicated minimal genetic differences between the populations with Least Chub. This resulted in smaller spring and wetland (Mock and Miller 2005; Mock and Bjerregaard 2007); however, systems and manmade ponds being the focus for establishing Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 Mock and Miller (2005) recommended that Least Chub be introduced populations (Figure 3). Landowner willingness, grouped into and managed within three geographic management potential threats, likelihood of success, impacts on the existing units (GMUs; Figure 2). The GMUs are managed independently native species community, and impacts on the source population to maintain potential adaptive differences that were not were factors that determined where Least Chub were introduced. detectable within molecular markers. Because potential habitats were the limiting factor, the translocation of extant populations did not have to occur in the GMU from which they originated; recipient habitats only needed The introduction of Least Chub into additional to be located in the Bonneville Basin and have no potential for locations began as early as 1979. Least Chub escapement into other extant populations or other private lands. One major action identified in the LCCAS was to establish and maintain three introduced Least Chub populations per GMU Establishing Introductions source with at least one per distinct extant population source for Utah Division of Wildlife Resources (UDWR) biologists a total of nine introduced populations (LCCT 2014). Though the used their expert opinions to determine suitable habitats, and term “introduced” can reference the stocking of nonnative fish or though this process was subjective, these general factors were the stocking of fish outside of their known range (Hendrickson considered: (1) stability of water source, (2) habitat protection and Brooks 1991; George et al. 2009), this was the terminology (e.g., minimal or no livestock grazing, remote, etc.), (3) presence utilized by the LCCT and the USFWS during the listing decision or diversity of fishes, and (4) size (e.g., small habitats are process, so throughout the remainder of this article, introduced more subject to catastrophic disturbances). Two habitats were

550 Fisheries | Vol. 40 • No. 11 • November 2015 Downloaded by [Department Of Fisheries] at 23:47 13 March 2016

Figure 3. Examples of habitats where Least Chub were introduced with SE Pilot (bottom) and Rosebud Pond (top). Photo credits: Paul Thompson.

Fisheries | www.fisheries.org 551 chemically treated with rotenone to remove nonnative fish had elapsed to determine whether Least Chub had become prior to the introduction of Least Chub. Other sites contained established at the six locations that were stocked from 2012 nonnative fish such as sterile Rainbow Trout Oncorhynchus to 2013. For the 17 locations that were evaluated, 10 of the mykiss, which were determined not to threaten Least Chub introduced populations met the introduced population criteria success, but the majority of introduction sites were free of and were considered successful introduced Least Chub nonnative fishes. Least Chub were collected with mesh minnow populations (Table 2; Figure 2). The three introductions into traps or seines from wild and hatchery populations. All hatchery lotic habitats were unsuccessful because Least Chub were not populations were created from wild-caught broodstock that were documented again at these sites (Table 2). The population at placed in earthen ponds or cement raceways with spawning Willow was considered a success for four years; however, a structure (e.g., submerged Christmas trees) and allowed to wildfire in 2013 and subsequent ash flow into the pond from reproduce and recruit in the absence of nonnative fishes and rain events severely reduced the number of Least Chub at this without human intervention. Least Chub were transported location. Least Chub were salvaged from this site following the in UDWR hatchery trucks or coolers with battery-powered fire and subsequent monitoring documented their persistence, aerators and, if needed, the fish were temperature acclimated to but as of 2014, Willow was not considered a successful the recipient waters prior to stocking. A minimum of 200 Least introduced population because post-fire threats remained Chub was chosen to establish new Least Chub populations with elevated. The Least Chub population at Stokes was considered a target of 500 fish stocked in larger habitats (e.g., >1 ha). If the unsuccessful because there was no formal agreement in place donor population was believed to be less than 1,000 individuals, with the private landowner. Annual Least Chub monitoring at transplants occurred over multiple years to minimize impacts to Chambers indicated that this population was small with minimal the donor population. recruitment, and there was no formal agreement with the private The UDWR followed Utah code and agency policies landowner. Shortly after the introduction of Least Chub into during the collection, transportation, and stocking of wild or Fish Springs National Wildlife Refuge, Western Mosquitofish hatchery Least Chub. Least Chub could only be moved if (1) invaded and this was attributed to the failure of this introduction. the donor population was certified pathogen free as defined in the American Fisheries Society Fish Health Section Blue Listing Decision Book (Mitchell and Goowin 2004), (2) the donor and recipient In the 2014 USFWS decision to remove Least Chub from the location were surveyed for aquatic invasive species to ensure candidate list, each of the 10 successful introduced populations there was no transport of invasive species with the fish to the were evaluated using the five factor threat assessment in recipient site, and (3) the introduction was supported. Support the same manner as the extant populations. The five factors for a Least Chub introduction was obtained from the LCCT and USFWS evaluates in every listing decision are as follows: (1) UDWR as well as the owner of the property and county where damage to or destruction of habitat; (2) overutilization of the the introduction would occur. In all cases the landowner was species for commercial, recreational, scientific, or educational made fully aware of the current federal candidate status for purposes; (3) disease or predation; (4) inadequacy of existing Least Chub. Initially, some private landowners were asked to protection; and (5) other natural or manmade factors. Based on sign an agreement with UDWR, establishing the conditions for this analysis in the 2010 decision that determined that Least the introduction and what both parties agreed to. One of these Chub were warranted for listing (USFSW 2010b), the threats stipulations allowed landowners to request UDWR to remove that were significant for Least Chub were as follows: factor Least Chub from their property with sufficient notice. A—specifically livestock grazing and groundwater withdrawal; factor C—introduction of nonnative fish; factor D—inadequacy Monitoring of existing regulatory mechanisms, specifically relating to A new population of Least Chub was considered a successful groundwater regulation; and factor E—the cumulative effects of introduced population if the population remained secure with drought and climate change. no or minimal threats (e.g., minimal livestock grazing, secure water source, and few or no nonnative fish present), there was Threats Analysis Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 a current agreement in place if the population was on private No significant threats were identified for the 10 introduced land, and the population had successfully spawned for at least Least Chub populations that were considered successful by the previous two years. Successful spawning was determined by the LCCT. Livestock grazing was completely excluded from the presence of age-0 Least Chub (<35 mm TL; modified from 6 of the 10 sites, and 4 allowed some level of grazing that was Mills et al. 2004b) during annual monitoring with mesh minnow not considered detrimental to Least Chub habitat. All of the traps or seines. Monitoring occurred in mid-August through introduced populations had a water right associated with the September and required the capture and measurement of 100 site. Though the water right was not exclusively for maintaining Least Chub per population and followed the same procedure Least Chub habitat, this provided a mechanism for ensuring as monitoring of the extant populations (LCCT 2010). The that water levels were maintained and that additional water population status for all introduced populations of Least Chub use in the future would not impact these habitats. With regard was reviewed annually by the LCCT to determine whether a to factor C, there were only two introduced populations that population should be considered successful. had any nonnative species present. These were Common Carp Cyprinus carpio and Goldfish Carassius auratus at Atherly RESULTS and sterile Rainbow Trout that were stocked at Rosebud Pond. Successful Introductions The introduced populations where these nonnative fish occur Between 2005 and 2013, Least Chub were introduced were two of the larger sites with complex habitat. Common into 23 locations (not including UDWR hatchery facilities Carp in large numbers can have a significant impact on habitat and educational facilities) within the Bonneville Basin, Utah (Parkos et al 2003; Matsuzuki et al. 2007), but they occurred (Table 2). As of the 2014 USFWS decision, insufficient time in low densities and were not negatively impacting the Least

552 Fisheries | Vol. 40 • No. 11 • November 2015 Table 2. Least Chub introductions in the Bonneville Basin, Utah, 2005–2013. Other species: UC = Utah Chub Gila atraria, MF = Western Mosquitofish, CP = Common Carp, GF = Goldfish, SD = Speckled Dace Rhinichthys osculus, RT = Rainbow Trout, BT = Brook Trout Salvelinus fontinalis, US = Utah Sucker Catostomus ardens, MT = Mountain Sucker C. platyrhynchus, and LB = Largemouth Bass Micropterus salmoides. NWR = National Wildlife Refuge, WD = West Desert, SR = Sevier River, WF = Wasatch Front. RI = recently introduced. Year Estimated Other Least Chub Population met Source popu- introduced Habitat water Site Ownership species recruitment introduced criteria of lation/GMU (# of Least type volume present documented LCCT (if not, reason) Chub) (m3) Red Knolls Bishop/WD 2005 Federal Pond 220 Nonea Yes Yes (250) Fish Springs Leland/WD 2006 Federal Marsh 1,466,610 MF, UC No No (not biologically NWR (170) successful) Atherly Mills Valley/SR 2006 State Reservoir 162,865 UC, CP, GF Yes Yes (19,000) Escalante Mona/WF 2006 Local Pond 52 None Yes Yes (175) government Willow Clear Lake/SR 2007 Private Pond 330 None Yes No (2013 wildfire (340) threatens population) Pilot Leland/WD 2008 Federal Pond 640 Nonea Yes Yes (147) 2013 (208) SE Pilot Mills Valley/SR 2008 Federal Pond 670 None Yes Yes (700) Rosebud Mills Valley/SR 2008 Private Reservoir 24,260 SD, RT Yes Yes Pond (3,000) Rosebud Mills Valley/SR 2008 Federal, Flowing n/a SD No No (not biologically Spring (500) private spring successful) Cluster Mills Valley/SR 2008 Federal Pond 62 None Yes Yes (two ponds) (400) Stokes Mills Valley/SR 2008 Private Pond 1,320 UC Yes No (no agreement with (700) landowner) Chambers Mills Valley/SR 2008 Private Pond 4,570 SD, CP Yes No (small population, (1,000) limited recruitment) Mountain Mills Valley/SR 2008 Private Flowing n/a SD, RT No No (not biologically Valley (1,200) spring successful) AW Mills Valley/SR 2008 Private Flowing n/a US No No (not biologically Heritage (1,700) spring successful) Keg Gandy/WD 2009 Federal Pond 570 None Yes Yes (200) Upper Mona/WF 2011 State Pond 385 None Yes Yes Garden (1,000) Deseret Mona/WF 2011 Federal Pond 978 None Yes Yes (500) Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 West Bishop/WD 2012 State Pond 28,330 MF, CP, UC No No (RI) Locomotive (341) 2013 (216) Sparks Mona/WF 2012 State Pond 7,280 MF, CP, UC No No (RI) (2,000) Teal Clear Lake/SR 2013 State Pond 60,700 MF, CP, No No (RI) (500) UC, LB Baker Gandy/WD 2013 State Pond 121,410 MF, CP, UC No No (RI) (496) Bar M Leland/WD 2013 State Pond 42,490 MF, CP, No No (RI) (605) UC, MT Big Springs Mona/WD 2013 Private Pond 8,250 BT No No (RI) (15,200)

aChemically treated with rotenone prior to the introduction of Least Chub.

Fisheries | www.fisheries.org 553 Chub populations. Sterile Rainbow Trout were stocked into one introduced populations located outside its historical range of the privately owned introduced populations as an incentive (Lintermans 2013a). for the landowner that allowed the UDWR to establish a Least The creation of self-sustaining introduced Least Chub Chub population on their property. This stocking was initially populations proved to be relatively simple, likely because they considered experimental; however, the habitat complexity at are generalists and have a broad tolerance limit to many water this site allowed sufficient habitat separation between Rainbow quality parameters (Sigler and Sigler 1987). Because Least Chub Trout and Least Chub and did not negatively impact the Least become reproductively mature in one year and are intermittent Chub population. Inadequacy of existing regulatory mechanisms spawners (Crawford 1979), releasing eggs throughout the also was not determined to be a threat at any of the introduced majority of late spring/summer months, populations often populations because there was often a water right associated became well established within two years from the introduction. with the site. All introduced populations were determined to be Least Chub that were introduced into smaller ponds in the managed by state or federal agencies or under an agreement with absence of other fish species generally became established. Of an agency. These agreements and the management authority the 10 successful introduced populations, Least Chub were the helped ensure that sufficient regulatory mechanisms were in only species present in eight of the habitats (Table 2). Although place at the introduced populations. Cumulative effects of Least Chub historically occupied lotic habitats (Sigler and drought and climate change were determined to be a threat in Sigler 1987; Page and Burr 1991), the three introductions into the 2010 decision for the extant Least Chub populations. The lotic habitats were unsuccessful (Table 2) for reasons unknown. introduced populations were a significant factor in addressing Hickman (1989) considered the Least Chub to be constantly this threat because they provided mitigation by providing threatened by the introduction and presence of nonnative fish redundancy and resiliency. species, and Western Mosquitofish pose a direct threat because of their known aggressive predation on eggs and young of other DISCUSSION fishes (Meffe 1985; Sigler and Sigler 1987). Mills et al. (2004b) Establishing introduced populations for sensitive fish species found that Least Chub juveniles were vulnerable to Western has been a common conservation practice. Such an approach Mosquitofish predation and no Least Chub introductions were frequently is considered when the species is endemic to a small considered successful when Western Mosquitofish were present range or has considerable threats to its existence in its current (Table 2). The invasion or illegal introduction of nonnative fish occupied range or simply as a precaution to avoid extinction species into extant and introduced Least Chub populations will from catastrophic events. Managers also need to consider the be a continual threat; however, the majority of these populations potential impacts of introducing the species on the native species occur in remote locations, which should lessen this threat. already existing in that location, as well as consider the impact The establishment of introduced populations is often an if the introduced species escapes or expands beyond its intended integral part of ESA recovery plans because range expansion location. Other factors to consider are the impacts of removing and establishment of new populations are typically a criterion fish on the source population. If the species is long-lived or slow for down-listing or delisting (USFWS 2000; Paragamian and to reproduce, it may take many years before the success of the Beamesderfer 2004); however, this goal is not always achieved. introduction can be determined; in addition, multiple transfers In Australia, no freshwater fish has had its conservation may be needed and this could impact the source populations status down-listed or delisted as a result of conservation (Minckley 1995; George et al. 2009). These factors were management, including the establishment of introduced considered in Least Chub introductions but were determined populations (Lintermans 2013b). Hendrickson and Brooks not to be significant barriers. Many of the introduced locations (1991) also provide many examples of imperiled fishes in the were artificially created ponds and any native species present western United States where introduced populations have been also coexisted with Least Chub in the extant populations. The successfully established, yet their official regulatory status was impact on the source populations was considered, but for two not changed as a result. In 2014, however, the Oregon Chub extant populations, a successful hatchery population already Oregonichthys crameri was removed from protection under ESA (USFWS 2014a). The Oregon Chub is similar to the Least Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 had been established and was utilized as a source of fish for the introductions. Chub in that it is a small minnow and shares some life history Introduced or conservation populations have been created similarities. The Oregon Chub Recovery Plan (USFWS 1998) for several species in the western United States, including the outlined the establishment of refuge populations to preserve Apache Trout Oncorhynchus apache, Big Bend Gambusia genetic variability among populations (DeHaan et al. 2012) Gambusia gaigei, Desert Pupfish Cyprinodon macularius, and was a requirement for delisting (USFWS 1998). The and Gila Topminnow Poeciliopsis occidentalis (Simons 1987; successful delisting of the Oregon Chub not only provides hope Johnson and Hubbs 1989; Williams 1991). Translocating fishes for the delisting of additional threatened or endangered fishes from the wild to new locations has saved some species from but highlights the important role that establishing introduced extinction, but establishing introduced populations for a species populations can serve in recovery. does not guarantee persistence (e.g., Amistad Gambusia G. The Least Chub was not a federally listed species under amistadensis; Hubbs and Jensen 1984). The establishment of ESA; however, in 2007, this fish was petitioned to be listed as introduced populations is not specific to the western United threatened or endangered under ESA including the designation States. In Australia, the Murray Hardyhead Craterocephalus of critical habitat. The USFWS concluded in a 90-day finding fluviatilis was taken into captivity and subsequently stocked that the petition did present substantial information in support into several locations with some success (Ellis et al. 2013). of listing (USFWS 2008). The 90-day finding resulted in a In addition, the Pedder Galaxias Galaxias pedderensis was 12-month status review that identified the Least Chub in 2010 found only in Lake Pedder in Tasmania, Australia. This species as a species for which listing as endangered or threatened was was removed from Lake Pedder and now exists only in two warranted but precluded (USFWS 2010a). The Least Chub remained a candidate species for four years until the USFWS

554 Fisheries | Vol. 40 • No. 11 • November 2015 ruled in 2014 that federal protection under ESA was not needed Mark Grover, Krissy Wilson, Paul Thompson, Aaron Webber, and the Least Chub was removed from the candidate list and Samuel McKay were the biologists and managers who (USFWS 2014b). worked diligently to find suitable habitats and introduce Least The 2014 decision cited that 10 successful Least Chub Chub. We thank Phaedra Budy and Mark Belk for reviewing introduced populations had been established and, when earlier drafts of this article. Reference to trade names does not combined with the naturally occurring populations, there was a imply endorsement by the U.S. Government. high likelihood of persistence even under higher probabilities of catastrophic events. The introduced populations were REFERENCES determined to provide habitat heterogeneity and redundancy Bailey, C. L., K. W. Wilson, and M. E. Andersen. 2005. Conservation and a buffer against environmental effects (e.g., cumulative agreement and strategy for Least Chub (Iotichthys phlegethontis) in the State of Utah. Utah Division of Wildlife Resources, effects from drought and changing climate conditions in the Publication No. 05-24, Salt Lake City. Bonneville Basin), and the geographic distribution of the Crawford, M. 1979. Reproductive modes of the Least Chub (Iotich- introduced populations was representative of the known genetic thys phlegethontis - Cope). Master’s thesis. Utah State Univer- diversity of the species (e.g., each natural population and sity, Logan. Deacon, J. E., G. Kobetich, J. D. Williams, and S. Contreras. 1979. GMU was represented in at least one introduced population). Fishes of North America endangered, threatened, or of special Therefore, if the Least Chub continued to persist in its current concern: 1979. Fisheries 4(2):29–44. distribution, there would be sufficient resiliency, redundancy, DeHaan, P. W., P. D. Scheerer, R. Rhew, and W. R. Ardren. 2012. Analy- ses of genetic variation in populations of Oregon Chub, a threat- and representation to persist now and in the foreseeable future ened floodplain minnow in a highly altered environment. Trans- (USFWS 2014a). actions of the American Fisheries Society 141:533–549. The introduced Least Chub populations that likely played Echelle, A. A. 1991. Conservation genetics and genetic diversity in the most significant role in the decision not to list Least Chub freshwater fishes of western North America. Pages 141–153 in W. L. Minckley and J. E. Deacon, editors. Battle against extinction were for the extant population at Mona (Least Chub were native fish management in the American West. The University of discovered in Mona Springs in 1995). The Mona population was Arizona Press, Tucson. not considered self-sustaining in the USFWS decision. Western Ellis, I. M., D. Stoessel, M. P. Hammer, S. D. Wedderburn, L. Suitor, and A. Hall. 2013. Conservation of an inauspicious endangered Mosquitofish and other nonnative fish also were documented freshwater fish, Murray Hardyhead (Craterocephalus fluviatilis), in 1995, and by 1998 the Least Chub population had declined during drought and competing water demands in the Mur- substantially. Least Chub were collected from Mona at that ray–Darling Basin, Australia Marine and Freshwater Research time and taken into captivity. Multiple efforts to remove 64:792–806. George, A. L., B. R. Kuhajda, J. D. Williams, M. A. Cantrell, P. L. Rakes, nonnative fish, improve habitat, and reestablish Least Chub were and J. R. Shute. 2009. Guidelines for propagation and transloca- attempted, but habitat complexity did not allow for the complete tion for freshwater fish conservation. Fisheries 34:529–545. removal of Western Mosquitofish. Recently, barriers have been Hendrickson, D. A., and J. E. Brooks. 1991. Transplanting short-lived fishes in North American deserts: review, assessment, and rec- established to create smaller habitats where nonnative removal ommendations. Pages 283–298 in W. L. Minckley and J. E. Dea- and Least Chub stocking efforts have been targeted. As of 2014, con, editors. Battle against extinction: native fish management Least Chub were successfully recruiting at Mona but they were in the American West. The University of Arizona Press, Tucson. still being stocked annually to maintain this population. Three Hickman, T. J. 1989. Status report of the Least Chub, Iotichthys phlegethontis, prepared for the U.S. Fish and Wildlife Service. introduced populations for the wild Mona population have been Western Ecosystems, St. George, Utah. established in addition to a hatchery population. The success and Hubbs, C., and B. L. Jensen. 1984. Extinction of Gambusia amis- stability of these introduced populations has ensured that any tadensis, an endangered fish. Copeia 1984:529–530. Johnson, J. E., and C. Hubbs. 1989. Status and conservation of south- unique genetic characteristic of the Mona Least Chub population western poeciliid fishes. Pages 301–317 in G. K. Meffe and F. F. will not be lost if a population cannot be maintained in the Snelson, Jr., editors. Evolution and ecology of livebearing fishes extant habitat. (Poeciliidae). Prentice-Hall, Englewood Cliffs, New Jersey. This was the first USFWS listing decision where introduced Lande, R. 1993. Risks of population extinction from demographic and environmental stochasticity and random catastrophes. fish populations were evaluated as discrete populations; hence, American Naturalist 142:911–927. the consideration of these introduced populations was a key LCCT (Least Chub Conservation Team). 2010. Least Chub monitor- Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 factor in precluding the need to list Least Chub under ESA. ing protocols. Utah Division of Wildlife Resources, Salt Lake City. ———. 2014. Amendment to the 2005 conservation agreement and Fisheries managers are faced with multiple threats to habitat strategy for the Least Chub (Iotichthys phlegethontis). Utah Di- that are difficult to address on site. Climate change receives vision of Wildlife Resources, Salt Lake City. consideration as a threat in most listing determinations and, Lintermans, M. 2013a. Conservation and management. Pages 283– consequently, the establishment of introduced populations 316 in P. Humphries and K. Walker, editors. The ecology of Aus- tralian freshwater fish. CSIRO Publishing, Melbourne, Australia. should be considered and may be one of the few tools for Lintermans, M. 2013b. A review of on-ground recovery actions for fisheries managers to prevent federal listings or recover listed threatened freshwater fish in Australia. Marine and Freshwater species. Research 64:775–791. Lydeard, C., and M. C. Belk. 1993. Management of indigenous species impacted by introduced mosquitofish: an experimental ap- ACKNOWLEDGMENTS proach. Southwestern Naturalist 38:370–373. We thank the highly functional Least Chub Conservation Matzusaki, S. S., N. Usio, N. Takamura, and I. Washitani. 2007. Effects of Common Carp on nutrient dynamics and littoral community Team for dedicating the past 16 years to providing solid composition: roles of excretion and bioturbation. Fundamental guidance for the conservation of the Least Chub and for helping and Applied Limnology 168:27–38. preclude the need for federal listing of this species under the Meffe, G. K. 1985. Predation and species replacement in American Southwestern fishes: a case study. Southwestern Naturalist ESA. The work to establish Least Chub by UDWR biologists 30:173–187. was funded by UDWR through Endangered Species Mitigation Miller, R. R. 1972. Threatened freshwater fishes of the United States. Funds and State Wildlife Grants, as well as the Bureau of Land Transactions of the American Fisheries Society 101:239–252. Management. Kevin Wheeler, Richard Fridell, Chris Crockett, Mills, M. D., M. C. Belk, R. B. Radar, and J. E. Brown. 2004a. Age and growth of Least Chub, Iotichthys phlegethontis, in wild popula-

Fisheries | www.fisheries.org 555 tions. Western North American Naturalist 64:409–412. ———. 2010b. Endangered and threatened wildlife and plants; review Mills, M. D., R. B. Rader, and M. C. Belk. 2004b. Complex interactions of native species that are candidates for listing as endangered between native and invasive fish: the simultaneous effects of or threatened; annual notice of findings on resubmitted peti- multiple negative interactions. Oecologia 141:713–721. tions; annual description of progress on listing actions. Federal Minckley, W. L. 1995. Translocation as a tool for conserving imperiled Register 75(217):69222–69294. fishes: experiences in Western United States. Biological Conser- ———. 2011. Endangered and threatened wildlife and plants; review of vation 72:297–309. native species that are candidates for listing as endangered or Minckley, W. L., and M. E. Douglas. 1991. Discovery and extinction of threatened; annual notice of findings on resubmitted petitions; western fishes: a blink of the eye in geologic time. Pages 7–17 in annual description of progress on listing actions. Federal Regis- W. L. Minckley and J. E. Deacon, editors. Battle against extinc- ter 76(207):66370–66439. tion: native fish management in the American West. The Univer- ———. 2012. Endangered and threatened wildlife and plants; review of sity of Arizona Press, Tucson. native species that are candidates for listing as endangered or Mitchell, A., and A. Goodwin. 2004. Centrocestiasis (gill trematode threatened; annual notice of findings on resubmitted petitions; disease). Pages in FHS blue book: suggested procedures for annual description of progress on listing actions. Federal Regis- the detection and identification of certain finfish and shellfish ter 77(225):69993–70060. pathogens, 2012 edition. American Fisheries Society Fish Health ———. 2013. Endangered and threatened wildlife and plants; review of Section, Bethesda, Maryland. native species that are candidates for listing as endangered or Mock, K. E., and L. S. Bjerregaard. 2007. Genetic analysis of a recent- threatened; annual notice of findings on resubmitted petitions; ly discovered population of the Least Chub (Iotichthys phleg- annual description of progress on listing actions. Federal Regis- ethontis). Western North American Naturalist 67:142–146. ter 78(226):70103–70162. Mock, K. E., and M. P. Miller. 2005. Patterns of molecular diversity in ———. 2014a. Endangered and threatened wildlife and plants; remov- naturally occurring and refugial populations of the Least Chub. ing the Oregon Chub from the list of endangered and threat- Transactions of the American Fisheries Society 134:267–278. ened wildlife. Federal Register 79(25):7136–7152. Moyle, P. B., and R. A. Leidy. 1992. Loss of biodiversity in aquatic ———. 2014b. Endangered and threatened wildlife and plants; ecosystems: evidence from fish faunas. Pages 127–169 in P. L. 12-month finding on the petition to list Least Chub as an endan- Fielder and S. K. Jain, editors. Conservation biology: the theory gered or threatened species. Federal Register 79(165):51041– and practice of nature conservation, preservation, and manage- 51066. ment. Chapman and Hall, New York. Williams, J. E. 1991. Preserves and refuges for native western fishes: Page, L. M., and B. M. Burr. 1991. A field guide to freshwater fishes of some protected sites and species, and recent federal legislation. North America north of Mexico. Houghton Mifflin, Boston. Pages 171–189 in W. L. Minckley and J. E. Deacon, editors. Bat- Paragamian, V. L., and R. C. P. Beamesderfer. 2004. Dilemma on the tle against extinction: native fish management in the American Kootenai River—the risk of extinction or when does the hatch- West. The University of Arizona Press, Tucson. ery become the best option? Pages 377–385 in M. Nickum, P. Williams, J. E., J. E. Johnson, D. A. Hendrickson, S. Contreras- Mazik, J. Nickum, and D. MacKinlay, editors. Propagated fish in Balderas, J. D. Williams, M. Navarro-Mendoza, D. E. McAllister, resource management. American Fisheries Society, Symposium and J. E. Deacon. 1989. Fishes of North America: endangered, 44, Bethesda, Maryland. threatened, or of special concern. Fisheries 14(6):2–20. Parkos, J. J., III, V. J. Santucci, and D. H. Wahl. 2003. Effects of adult Common Carp (Cyprinus carpio) on multiple trophic levels in shallow mesocosms. Canadian Journal of Fisheries and Aquatic Sciences 60:182–192. Perkins, M. J., L. D. Lentsch, and J. Mizzi. 1998. Conservation agreement and strategy for Least Chub (Iotichthys phlegethontis) in the State of Utah. Utah Division of Wildlife Resources, Publica- tion No. 98-25, Salt Lake City. Peterson, J. T., and J. Sanez. 2013. The development of an adaptive, decision-support tool for the conservation and recovery of Least Chub, Iotichthys Phlegethontis. Draft interim final report to the Least Chub Conservation Team. Sigler, W. F., and J.W. Sigler. 1987. Fishes of the Great Basin, a natural history. University of Nevada Press, Reno. ———. 1996. Fishes of Utah a natural history. University of Utah Press, Salt Lake City. Simons, L. H. 1987. Status of the endangered Gila Topminnow, Po- eciliopsis occidentalis, in the United States. Arizona Game and Fish Department, Project E-I, Title VI, Phoenix. Stevens, L. E., and V. J. Meretsky. 2008. Aridland springs in North

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 America. Ecology and conservation. University of Arizona Press, Tucson. USFWS (U.S. Fish and Wildlife Service). 1982. Endangered and threatened wildlife and plants; review of vertebrate wildlife for listing as endangered or threatened species. Federal Register 47(351):58454–58460. ———. 1989. Endangered and threatened wildlife and plants; animal notice of review. Federal Register 54(4):554–579. ———. 1995. Endangered and threatened wildlife and plants; proposal to determine the Least Chub (Iotichthys phlegethontis) an endangered species with critical habitat. Federal Register 60(189):50518–50530. ———. 1998. Recovery plan for the Oregon Chub (Oregonichthys crameri). Portland, Oregon. ———. 2000. USFWS–NMFS policy regarding controlled propagation of species listed under the ESA. Federal Register 65:56916– 56922. ———. 2008. Endangered and threatened wildlife and plants; 90-day finding on a petition to list the Least Chub (Iotichthys phlegethontis) as threatened or endangered with critical habitat. Fed- eral Register 73(200):61007–61015. ———. 2010a. Endangered and threatened wildlife and plants; 12-month finding on a petition to list the Least Chub as threatened or endangered. Federal Register 75(119):35398–35424.

556 Fisheries | Vol. 40 • No. 11 • November 2015 ESSAY Relevant Topics to Keep in Mind as a Graduate Student Downloaded by [Department Of Fisheries] at 23:47 13 March 2016

Artur Rombenso Center for Fisheries, Aquaculture, and Aquatic Sciences and Departments of Zoology and Animal Science, Food and Nutrition, Southern Illinois University Carbondale, 1125 Lincoln Drive Room 251, Carbondale, IL 62901. E-mail: [email protected]

Karma Kissinger Tropical Conservation Biology and Environmental Science, University of Hawaii at Hilo, Hilo, HI

Michael Ciaramella Department of Food Science, Nutrition and Health Promotion and Wildlife, Fisheries and Aquaculture, Mississippi State University, Mississippi State, MS

Jesse Trushenski Center for Fisheries, Aquaculture, and Aquatic Sciences and Departments of Zoology and Animal Science, Food and Nutrition, Southern Illinois University Carbondale, Carbondale, IL

Fisheries | www.fisheries.org 557 INTRODUCTION MAKING THE RIGHT DECISION Job search and employment have never been as competitive Before making the big decision about whether or not to go as they are today. Globalization, high-speed information to grad school, it is essential to know (or at least have an idea exchange, increased global population and thus numbers of of) what you want to do in future, what kind of job you will job-seekers, and global economic crises are some of the major look for, and what your dream job requires. This may be the reasons for the job market’s slowdown. As a result, opting most important thing to figure out before searching for the best for a higher education has been considered a positive strategy grad school because all jobs require potential candidates to have to be successful in the job hunting process. In addition to the distinct skillsets and qualifications; for instance, some jobs in monetary gratification provided by a graduate degree, there are fisheries and aquaculture do not require a graduate degree but other relevant achievements such as personal growth, credibility, instead require more practical field and work experience. An specialization in your area of interest, career advancement, and aquaculture technician position in a hatchery or a production more employment opportunities. However, life as a graduate farm will require extensive practical experience working student is very different from that of the undergraduate outdoors, maintaining different culture systems, monitoring experience, and the transition can be challenging, especially water quality, and maybe even assessing disease. Other skills when there is a lack of knowledge regarding expectations including good written and oral communication, the ability to of grad students and uncertainty as to whether or not grad work alone or in a team, and a research background to be able to school is right for you. Few students have a clear idea of what assist in possible research trials might also be requested. These to expect from higher education and are often overwhelmed needs are quite different from a position that requires a graduate with a new schedule full of responsibilities, including classes, degree—for example, a hatchery researcher or manager, which research, teaching, writing manuscripts, giving presentations, usually requires a master’s or other advanced degree. and organizing events, among others miscellaneous tasks. There are several ways to get more familiar with the Without good orientation, planning, organization, and support, requirements and skillsets necessary for your ideal job. Online the pressures of grad school can be quite stressful. For some, searching can provide a general idea of what most employers the stress experienced during the early stages of graduate require, talking with your supervisor and others in your school may incite a desire to quit the program—even students department will give you important feedback, and contacting who would otherwise do quite well after learning the ropes can specialists in your area of interest and contacting current feel this way. Graduate education should not be an unpleasant workers to receive a real perspective on their daily routine and experience; rather, it should be an enjoyable period of one’s life. tasks can give you great insight. Though those recommendations seem obvious, they are easy to overlook, so it’s worth double- checking to make sure you are on the right path before jumping Before making any decisions about grad into something that may not be useful to you down the road. Grad school is a lot of work, and you want to make sure that it school, you need to know what educational, serves a purpose. professional, and practical requirements Once you’ve decided that graduate school is your next step, there are a few things you should consider in order to make there are for your ideal career. yourself competitive and likely to secure a place in the graduate program of your choice. Things prospective advisors may look for on an application or during an interview include the As the saying goes, failing to plan is planning to fail. Before following: making any decisions about grad school, you need to know what educational, professional, and practical requirements 1. How much experience does the applicant have and does it there are for your ideal career. This seems obvious, but students seem legitimate? What was the nature of their experience sometimes pursue graduate school without necessity—some and is it relevant to this position/program? Is the applicant’s

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 careers don’t require advanced degrees. In situations like this, experience relevant to the position/program and does his or practical work experience and specialized courses are more her educational background indicate that he or she would valuable than a graduate degree. So knowing where you want to be adequately prepared for graduate school? Sell yourself, work and what that requires is an essential step before jumping but don’t pad your CV! Advisors can usually tell if a student into grad school. Once you’ve determined that graduate school is exaggerating his experience in his application; they will is an important step on your career path and you’ve identified a definitely know what your strengths and challenges are after program of study, some additional planning and preparation can they begin supervising you. help make the transition and graduate experience much easier 2. Is there evidence of good writing skills? Does the application and more enjoyable. If everything is well organized and one show clear, concise writing, and is it free of typos and enters a program with realistic expectations, graduate school can grammatical errors? Has the student published, presented, or be a rewarding and successful experience. written successful grant proposals before? To help you on your journey, we have put together the 3. Does the applicant use good communication skills, articulate following guidelines to help undergraduates and graduate ideas well, and use concise speech? A phone or in-person students (current or prospective) navigate the decision-making interview can seal the deal or break it. Applicants who and preparation processes. These guidelines were compiled from approach interviews well prepared and professionally and several surveys of former grad students (currently employed) have done their homework about the program/position tend and employers. to perform better than those who don’t.

558 Fisheries | Vol. 40 • No. 11 • November 2015 Networking isn’t just about meeting people but keeping in touch with those you meet.

4. Character/attitude/personality. Is the applicant personable? • In addition to research experience with fish or other Will she work well with others in the lab? Does she organisms, it will be valuable to have skills pertinent to demonstrate initiative and a good work ethic? the job (i.e., microbiology, hands-on skills with carpentry/ 5. Grades and GRE scores matter, but so do experience, letters plumbing/electrical, and maybe small engine repair, know of recommendation, and attitude. Not all successful people how to build systems, use power tools, calculate flow rates, perform well in the context of standardized testing, and calculate tank values, apply basic geometry and algebra in most advisors understand this. Though poor grades or GRE problem-solving situations, etc.). scores do not strengthen an application, applicants with the right attitude and other evidence of scholarly promise can Communication still be successful in securing graduate positions and earning Effective communication, both written and oral, will be advanced degrees. integral to most jobs you encounter in the future, especially if you are looking for a higher-level position. Take time during GETTING THE MOST OUT OF YOUR your studies to learn to write succinctly and concisely. Here are GRADUATE EXPERIENCE a few ways you can work on your communication skills: Most students cannot wait to be done with college or grad school and move out into the “real world” with a real job, where • Familiarize yourself with scientific and professional they no longer have to worry about classes, tests, and other documents. Find and read environmental assessments, student responsibilities. However, being a student is a unique environmental impact statements, requests for proposals, and moment that passes quite fast. Although the responsibilities successfully funded grant proposals.

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 of a graduate student may seem daunting at times—balancing • Whenever possible, publish your work. For some careers, teaching, research, attending meetings, writing reports, meeting publications are not a requirement, and not everyone enjoys deadlines, and supervising student workers—responsibilities and writing and seeing their name in print. That said, publishing the challenges of time management will only increase as you your work is an important part of the graduate experience, advance in your career. So enjoy this period as much as possible and publishing your work in the peer-reviewed literature is and be open to all opportunities. Take advantage of your ability the gold standard of scientific communication. Publications to gain these experiences while you are a student, because they are extremely important for those seeking employment in will undoubtedly help you advance further and better equip you academia, but they strengthen your CV/résumé for positions for a successful career with a graduate degree. with state and federal agencies, industry, and commercial The following is a list of 10 areas to focus on to ensure that ventures, too. So work hard and try to publish as much as you get the most out of your time (Table 1). you can. Remember that magazines, blogs, and other social media can be an effective means of promoting your work Professional Experience and letting people know what you are doing! As much as In addition to completing your degree requirements possible, publish prior to graduating; it is very difficult to get and thesis/dissertation research, work or volunteer at a things completed once you get out of school. Jobs, family, commercial farm or another research facility. Try to do this life in general tends to take over, and you don’t always early in your school career so you can choose your classes based have access to electronic databases and statistical analysis on interest and get hands-on experience! software (or the day-to-day help of your advisor) after graduation.

Fisheries | www.fisheries.org 559 keeping in touch with those you meet. Work toward developing good relationships, meeting new people, and making meaningful connections because this may be crucial in achieving your future goals. Take advantage of conferences, seminars, and other gatherings at your university and be an active member of professional organizations—each of these provides ample opportunities to make new connections. Talk to your supervisor and other professors in your department because they likely know key people and organizations that may be of interest to you. In fisheries and aquaculture, everyone knows everyone, and this will help when it comes time to applying for jobs and supplying references to potential employers. • Get letters of reference from professors lined up well before graduation. Make sure that your professors give you a good verbal recommendation, too. Employers who are looking to hire often go to aquaculture professors at a few select schools whose programs they trust.

Leadership Experience With a higher education, the jobs you seek will most Showing initiative and volunteering your time likely require you to lead or manage one employee or shows commitment and will help you gain several others on a regular basis. Getting experience through holding leadership positions and advising technicians and valuable contacts and references. undergraduates is a great way to show potential employers that you are capable of doing that. Taking a leadership position in • Attend and present at as many conferences as possible. the graduate student association or other graduate student clubs Hopefully, you will have an opportunity to give a standard can be a good way to gain such experience. Hiring technicians 10- to 20-minute technical presentation during your time as a to help you conduct your research gives you valuable hands- student. It’s also useful to create a poster or two and practice on experience with mentoring, advising, and teaching. Many giving a quick five-minute synopsis (an elevator speech) programs will already have funding to hire technicians to help and longer lectures. Conferences are great opportunities with various aspects of research projects, but if yours does to network, share your research and ideas, and meet new not, you can write a small grant proposal to support hiring a people. So attend as many meetings/conferences/workshops technician and develop your writing skills in the process! There as you can. are many sources of funding available that could be used for this • Apply for awards, grants, and fellowships to support purpose. your research and attend national and international conferences. There are so many opportunities for research Organization and travel/conference awards—you just need to look for Being organized and timely will be integral to your them. Many institutions offer support and awards to current success in grad school and in your future career. As you students, as do private foundations, industry coalitions, advance in your education and professional career, you will professional organizations, and public agencies. Be positive gain more responsibilities and have more and more tasks to fit and apply for any and every one for which you are eligible— into your busy schedule. There will always be deadlines and timelines that must be met, and getting a head start on managing

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 receiving awards provides monetary benefit but also builds your CV/résumé. You may not get them all, but each one your time and resources effectively could make or break your will help improve your writing skills and give you a better graduate or professional education. Be sure to think ahead understanding of what committees looks for in applicants. because time will fly by and deadlines and opportunities will • Don’t forget there is always room for improvement, so pass you by in the blink of an eye if you are not careful. Here are take advantage of your program and improve on the a few tips to keep in mind. skills you already have. To become a well-recognized researcher/professor/manager you need to be able to speak • Plan and schedule your daily, weekly, and monthly tasks. to different audiences clearly and concisely; be able to Be knowledgeable of specific deadlines set in place by your communicate effectively in the form of reports, papers, department or university. Ensure that paperwork is prepared and other media; and provide leadership and mentoring to well in advance to allow ample time for processing and those around you. Spend some time enhancing the skillsets review by your advisor, committee members, and university you already possess and recognize there is always room for officials. improvement. Grad school is a great environment in which • Prepare/think about your next step. Do you know what to get constructive criticism and help from many other to do after graduation? No? Don’t worry, but you should professionals. spend some time strategizing your next steps. You should at least have some idea in mind of a general direction or area of Networking interest that you want to pursue. Start searching for jobs and • Make sure to network with those in the field and at graduate positions that appeal to you. This can help guide school. Networking isn’t just about meeting people but you in making this decision.

560 Fisheries | Vol. 40 • No. 11 • November 2015 Table 1. Top 10 things to keep in mind as a graduate During the graduate program we are busy, but taking student. advantage of travel opportunities can be a good way to recharge 1. Get professional experience (internships) your batteries, visit a new place, volunteer and gain additional skills, etc. After graduate school, opportunities to travel may be 2. Develop your skills (writing and speech) fewer and obligations related to your job, family, etc., may make 3. Network it more challenging to take advantage these opportunities. Don’t 4. Get leadership experiences be afraid to get out of your comfort zone. Go explore and gain 5. Be organized new experiences, both professional and personal.

6. Get involved and be an active member of a professional Collaboration organization or other community Work on side projects (outside of your thesis/dissertation 7. Get practical experience (attend specialized training) research) and be open-minded and ready to learn in order to 8. Be collaborative (participate in side projects) expand your knowledge base. This can include helping other 9. Get education & teaching experiences students/professors with their research projects or taking the lead on a project outside your comfort zone, working with others 10. Find a program you can be passionate about more familiar with that field. Be proactive; don’t think small. Diversifying your experiences will make you more marketable • Find out what you need to be prepared (example: and capable in the long run. develop a good CV or résumé, practice a second Education/Teaching language, etc.). As mentioned earlier, being able to effectively convey • Work on your CV/résumé and practice interviewing. your research to a variety of audiences is integral to • Find employers or organizations that focus on your achieving success in your future career. The best way you specialization, etc. can better prepare yourself for this is to teach. Professors spend a good portion of each semester preparing, presenting, and • Start applying for jobs, Ph.D. graduate programs, post- grading for the classes they teach. Those graduate students who doc positions (whatever your next step will be) at least serve as teaching assistants know how much time is invested six months before you graduate. Securing a position often in and outside the classroom. Offering to cover a lecture or two takes time, not only because you might not get the first for your advisor will make his week—maybe even their month! one you apply for, but because the application and review It saves him time and allows you to get practical experience process is often time consuming. If applying for a Ph.D. or teaching others. This will also be a good opportunity to receive post-doc position, try to define your project before entering constructive feedback on your teaching methods. Other ways to the program because this can save you time and avoid get teaching experience include volunteering to work summer inconveniences. The World Aquaculture Society (was.org), camps and workshops that many universities host. American Fisheries Society (fisheries.org), and USA Jobs (usajobs.gov) websites are all good places to look for jobs Passion and educational opportunities. Choose an advisor whose mentoring style meshes well with your needs as a student and a thesis/dissertation that Involvement/Commitment you can be passionate about. No two advisors or graduate Get involved. Your first priority should be to fulfill your students are exactly the same, and it’s important to make sure responsibilities to your advisor and the graduate program, but that you discuss your expectations/needs with your advisor in the remaining time, join clubs and volunteer to help with and clearly understand her expectations/needs. Initiating and other students on their projects. Helping with other students’ and maintaining an active line of communication with your advisor professors’ research will give you valuable, diverse, hands-on (or prospective advisor) doesn’t guarantee an easy path to

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 experience that could set you apart from other job-seekers/ success, but it does guarantee that problems will be identified applicants in the future. Showing initiative and volunteering early when it’s easiest to address them. It isn’t always possible your time shows commitment and will help you gain valuable to do exactly what you want for your graduate work, but you contacts and references. should pursue an advisor and project that you value and can enjoy. If your project is not compelling to you, it will be that Practical Experience/Specialized Training much more difficult to put in the time and hard work needed to Take advantage of all opportunities that come your way, successfully complete your degree. so participate, collaborate, and attend specialized courses and workshops. Look for workshops and certification classes in CONCLUSIONS your area of interest and get as much experience as you can! Try to find time for specialized trainings that provide credible The decision to attend graduate school is not an easy certifications. Some examples include the following: one, and advanced degrees are not easily earned. Graduate • Boating and electrofishing safety, usage of statistical and students experience stress but can also enjoy camaraderie, modeling software (e.g., SAS, R, GIS), commercial diving, fellowship, professional development, mentoring, and the etc. unique and powerful experience of learning through research • Take a few business courses to understand a basic business and the creation of new knowledge. The tips and considerations plan! presented here are by no means a comprehensive list but will hopefully help you navigate these sometimes murky waters with greater confidence and success.

Fisheries | www.fisheries.org 561 ESSAY Enhancing the Utility of the NHDPlus River Coverage: Characterizing Ecological River Reaches for Improved Management and Summary of Information Daniel J. Wieferich Department of Fisheries and Wildlife, Michigan State University, 1405 S. Harrison Rd STE 318b, East Lansing, MI 48823. E-mail: [email protected]

Wesley M. Daniel and Dana M. Infante Department of Fisheries and Wildlife, Michigan State University, East Lansing, MI Downloaded by [Department Of Fisheries] at 23:47 13 March 2016

The National Hydrography Dataset Plus (NHDPlus) is a publicly available geospatial hydrologic framework data set widely used as a tool for researching, modeling, and assessing characteristics of lentic and lotic freshwater systems. For streams, the smallest spatial unit of the NHDPlus, known as a “flowline,” mostly represents stream reaches formed by ecologically distinct hydrologic features of the geospatial framework. However, the NHDPlus also contains nonecological flowlines that are not defined by ecological criteria but are instead artifacts of cartography. We identified 129,773 nonecological flowlines in the NHDPlusV1 and 133,111 nonecological flowlines in the NHDPlusV2. We then developed a table (EcoReach) that assigns all flowlines into characterized ecological stream reaches. The publicly available table will facilitate summarization and reporting of results specific to ecological river reaches without altering the original version of the NHDPlus V1 or V2 and therefore allows for transferability of information with current and past research using the NHDPlus V1 or V2, respectively.

562 Fisheries | Vol. 40 • No. 11 • November 2015 REPRESENTATION OF STREAM REACHES IN THE NATIONAL HYDROGRAPH DATASET PLUS The National Hydrography Dataset Plus (NHDPlus; www. horizon-systems.com/NHDPlus/index.php) is a publicly available geospatial hydrologic framework data set widely used by agencies and researchers as a tool for researching, modeling, and assessing characteristics of freshwater systems, including rivers. The NHDPlus is available throughout the conterminous United States and is composed of multiple spatial units, with a basic unit defined by 1:100,000-scale flowlines representing unique segments of streams within fluvial networks (McKay et al. 2010, 2012). Flowlines have associated catchments also represented in the NHDPlus, with boundaries derived from digital elevation models. These spatial units can be used to identify or summarize characteristics of individual or multiple flowlines and their catchments, an essential step in developing stream reach–specific summaries of information, especially over large regions. Most flowlines within the NHDPlus data set represent stream reaches formed by ecologically distinct hydrologic features of the geospatial framework. These “ecological stream reaches” may be defined by stream origins, stream confluences, lake or reservoir inflows or outflows, and/or stream outflows (described in more detail in Wang et al. 2011). However, the NHDPlus also contains flowlines that have not been defined by ecological criteria but are instead artifacts of cartography. Within the NHDPlus data set, flowlines cross map boundaries, including U.S. Geological Survey (USGS) topological quadrangle maps and digital orthophoto quandrangle maps. At most intersections between flowlines and map boundaries, ecological stream reaches are split into multiple flowlines, resulting in development of “nonecological stream reaches.” This artifact of the data set has resulted in cases where ecological stream reaches are actually represented by multiple flowlines in the data set (Figure 1). Though these nonecological flowlines serve an important purpose in fully characterizing flow pathways throughout river networks, use of these units for attribution or summary of key pieces of information may confound the Figure 1. Stream network intersecting a USGS quadrangle boundary, summary of project results. where comid is a common identifier for flowlines and associated catchments. Ecological breaks occur at confluences, origins, and IDENTIFYING AND MANAGING FOR outflows of streams, whereas nonecological breaks occur at intersections of flowlines and USGS quadrangle boundaries. NONECOLOGICAL STREAM REACHES Using a combination of tabular and spatial data queries, The EcoReach table assigns an ecological grouping id Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 we identified 129,773 flowlines in the hydrological framework (ecoid) to every NHDPlus flowline and its associated catchment of the NHDPlusV1 (Figure 2) and 133,111 flowlines in the using the "comid" field, which is a common identifier assigned NHDPlusV2 that were created due to breaks in flowlines in the NHDPlus shared by flowlines and associated catchments. resulting from crossing quadrangle map boundaries. We found Steps for using this table to perform management or summary instances of this occurring in all 18 processing units of both of information for ecological stream reaches may differ the NHDPlusV1 and NHDPlusV2 data sets. To address this we depending on user preferences and needed information, but the developed a table (EcoReach) that assigns all flowlines into following steps are recommended: (1) attribute data to each of characterized ecological stream reaches (Wieferich et al. 2014a, the NHDPlus flowlines and/or associated catchments, (2) join 2014b). This EcoReach table and detailed metadata describing the EcoReach table to the summaries using the comid field as processing steps are publicly available for download on at the the join field (Table 1), (3) aggregate summary data to ecoids, following website: (ecosystems.usgs.gov/fishhabitat/nfhap_ (4) use the data associated with ecoids to perform analyses, and download.jsp). Following the processing steps described in the (5) link results back to the comids using EcoReach tables for metadata, users can also create similar EcoReach tables to group mapping and/or summary. nonecological flowlines resulting from flowline intersection This tabular identification of nonecological flowlines with quadrangle boundaries into ecological river reaches for resulting from flowline intersection with quadrangle boundaries other versions of the National Hydrography Datasets (e.g., NHD aids in characterizing information from the NHDPlus 1:24,000 in the conterminous United States, versions of the specifically for ecologically defined units. This product does not NHD in Hawaii and Alaska) as well as in future versions of the alter the original version of the NHDPlus V1 or V2 and therefore NHDPlus. allows for transferability of quantified and summarized variables

Fisheries | www.fisheries.org 563 Figure 2. NHDPlusV1 flowlines that are nonecological (n = 129,773), occurring where the NHDPlus stream coverage intersects boundaries of USGS quadrangle maps. Table 1. Using the EcoReach table with the stream network shown in Figure 2 as a reference. The Attributed comid column shows length and area attributed to each NHDPlus flowline and associated catchment, where comids 1, 2, 3, 4 are nonecological river reaches and 5, 6 are ecological river reaches. The EcoReach column, in the center, assigns ecological groupings (i.e., ecoid) to comids. The Attributed comid column is linked to the EcoReach column by the common field (comid) to allow for the summarization of attributes into ecoids

shown in the columns on the right, Attributed ecoid. In this example, lengths of flowlines including an ecoid are summed into lengthe in the Attributed ecoid column. Areas of associated catchments are summarized in the same manner. Attributed comid EcoReach Attributed ecoid

2 2 Comid length (km) area (km ) Comid Ecoid Ecoid Elength (km) Earea (km ) 1 3.25 10.00 1 1 1 6.25 10.50 3 3.00 5.50 3 1 2 1.75 6.00 2 2 2 5.25 11.00 4 3.50 5.00 4 2 5 1.20 3.00 5 3 3 1.20 3.00 6 1.25 2.00 6 4 4 1.25 2.00 Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 with current and past research using the NHDPlus V1 or V2, REFERENCES respectively. Ultimately, this table will facilitate the work of McKay, L., T. Bondelid, T. Dewald, C. Johnston, R. Moore, and A. Rea. researchers and modelers wishing to summarize or report results 2012. NHDPlus Version 2: user guide. Unites States Environmen- specific to ecological river reaches and/or their catchments. tal Protection Agency. Available: www.horizon-systems.com/ NHDPlus/index.php. (October 2014). McKay, L., T. Bondelid, C. Johnston, R. Moore, and A. Rea. 2010. ACKNOWLEDGMENT NHDPlus Version 1 (NHDPlusV1): user guide. United States En- We acknowledge Michael Slattery and Drew Ignizio for the vironmental Protection Agency. Available: www.horizon-systems.com/NHDPlus/NHDPlusV1_documentation.php. (October review of the data product. 2014). Wang, L. Z., D. Infante, P. Esselman, A. Cooper, D. Wu, W. Taylor, FUNDING D. Beard, G. Whelan, and A. Ostroff. 2011. A hierarchical spatial framework and database for the national river fish habitat condi- This product was developed with support from the U.S. tion assessment. Fisheries 369:436–449. Geological Survey Aquatic GAP Analysis Program and was Wieferich, D., W. M. Daniel, and D. M. Infante. 2014a. Ecological reach also supported by the U.S. Fish and Wildlife Service as part of identification table for NHDPlusV1: Version 2.0. National Fish Habitat Partnership Data System, U.S. Geological Survey, Res- the national inland assessment being conducted for the National ton, Virginia Fish Habitat Partnership. ———. 2014b. Ecological reach identification table for NHDPlusV2: Version 2.0. National Fish Habitat Partnership Data System, U.S. Geological Survey, Reston, Virginia

564 Fisheries | Vol. 40 • No. 11 • November 2015 2016 AFS ANNUAL MEETING 146th Annual Meeting of the American Fisheries Society: Third Call for Papers

It is not too soon to begin thinking about the 2016 Annual Meeting of the American Fisheries Society in Kansas City, Missouri, to be held August 21-25 at the Sheraton Crown Center. The conference will culminate President Essig's year- long message of "Fisheries Conservation and Management: Making Connections and Building Partnerships." While big river ecology and inter-jurisdictional fisheries will be a focus of several technical sessions, a diverse array of topics from marine fisheries to socio-economics will be highlighted at the conference. Start thinking now about how you can make a connection and build upon partnerships in Kansas City.

PROGRAM COMMITTEE CONTACTS Program Co-chairs: Country Club Plaza holiday lights. Abigail Franklin Archer Photo credit: VisitKC. Cape Cod Cooperative Extension Tel. 508-375-6702 E-mail: [email protected] SYMPOSIA Quinton Phelps • Proposals for Symposia must be submitted by January 17, Missouri Department of Conservation 2016. Tel. 573-243-2659 ext 1049 • Accepted symposia proposals will be posted on the E-mail: [email protected] conference website on February 15, 2016. • If accepted, organizers must submit a complete list of Contributed Papers Subcommittee Chairs: confirmed presentations and titles by March 6, 2016. Paul Michaletz Missouri Department of Conservation • Abstracts for symposium oral presentations must be Tel. 573-815-7901 ext 3921 submitted by March 13, 2016. E-mail: [email protected] Cari-Anne Hayer CONTRIBUTED PAPERS AND POSTERS USGS - Columbia Environmental Research Center • Those who wish to present in Contributed Papers or Poster Tel. 573-875-5399 sessions at the 2016 AFS meeting are required to submit E-mail: [email protected] Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 abstracts by February 14, 2016. This includes Student Symposia Subcommittee Chairs: Presentations. Sara Tripp • Confirmation of acceptance or refusal of abstracts will Missouri Department of Conservation be communicated by April 18, 2016. (Student presentations Tel. 573-243-2659 ext 1041 will be considered for a “best presentation” award if the E-mail: [email protected] student fills out additional application paperwork available at Heather Garrison www.fisheries society.org/education/BSP.htm.) Missouri Department of Conservation Tel. 573-468-3335 E-mail: [email protected] FOR MORE INFORMATION: VISIT: Craig Paukert 2016.fisheries.org Missouri Cooperative F&W Research Unit • AFS does not waive registration fees for presenters at Tel. 573-882-3524 symposia or contributed papers sessions or workshops. E-mail: [email protected] Registration forms will be available on the AFS website Poster Subcommittee Chair: (2016.fisheries.org) in May 2016; register early for cost Joanna Whittier savings. University of Missouri Tel. 573-884-7553 E-mail: [email protected]

Fisheries | www.fisheries.org 565 Journal Highlights NORTH AMERICAN JOURNAL OF AQUACULTURE Volume 77, Number 4, October 2015

Growth Performance of Spotted Rose Snapper in Floating Feeding Frequency and Rate Effects on Growth and Cages and Continuous Water-Flow Tank Systems. C. Physiology of Juvenile Genetically Improved Farmed Nile Hernández, L. Ibarra-Castro, C. H. Hernández, G. Quintero- Tilapia. Qing Huang, Kai Huang, Yanqun Ma, Xi Qin, Yanhong Martínez, E. A. Aragón-Noriega, and A. G. Tacon. 77:423-428. Wen, Lei Sun, and Lining Tang. 77:503-512.

[Communication] The Effect of a Mycotoxin Deactivation Propagation of Hybrid Devils Hole Pupfish × Ash Meadows Product on Growth of Juvenile Rainbow Trout Fed Distillers Amargosa Pupfish.Olin G. Feuerbacher, Justin A. Mapula, and Dried Grains. Wendy M. Sealey, Christopher G. Hooley, Kurt Scott A. Bonar. 77:513-523. A. Rosentrater, T. Gibson Gaylord, and Frederic T. Barrows. 77:429-436. Considerations for Consistently Applying Flow-Through Chloramine-T Treatments to Hatchery Raceways. James D. Effects of Stocking Density on Production Traits of Channel Bowker, Daniel G. Carty, Jesse T. Trushenski, David C. Glover, Catfish × Blue Catfish Hybrids. Brian Bosworth, Brian Ott, and Molly P. Bowman. 77:524-531. and Les Torrans. 77: 437-443. [Communication] Induced Spawning and Larval Culture [Technical Note] Effects of Rearing Density on Total Length of Golden Trevally. Jason S. Broach, Cortney L. Ohs, Andrew and Survival of Lake Sturgeon Free Embryos. John M. Palau, Bryan Danson, and Daniel Elefante. 77:532-538. Bauman, Edward A. Baker, Terry L. Marsh, and Kim T. Scribner. 77:444-448. Use of Lipid-Extracted Distillers Dried Grain with Solubles (DDGS) in Diets for Pacific White Shrimp. Melanie A. [Technical Note] Feeding and Growth of Larval Pacific Rhodes, Denghang Yu, Yangen Zhou, and D. Allen Davis. Lamprey Reared in Captivity. Jeffrey C. Jolley, Christina T. 77:539-546. Uh, Gregory S. Silver, and Timothy A. Whitesel. 77:449-459. [Technical Note] Adult Survival of Hatchery Spring Chinook [Communication] Research Verification of Production Salmon Released Volitionally or Forcibly as Juveniles. Practices Using Intensive Aeration at a Hybrid Catfish McLain S. Johnson, Andrew R. Murdoch, and Christopher P. H. Operation. Lisa B. Bott, Luke A. Roy, Terrill R. Hanson, Jesse Moran. 77:547-550. Chappell, and Gregory N. Whitis. 77:460-470. Referee Acknowledgments and Author Index for Volume 77 [Technical Note] Optimizing Time of Initiation for Triploid Walleye Production Using Pressure Shock Treatment. Eric R. Fetherman, Jesse M. Lepak, Bonnie L. Brown, and David J. Harris. 77:471-477.

Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 Tolerance and Avoidance Behavior towards Gas Supersaturation in Rock Carp Procypris rabaudi with a History of Previous Exposure. Yuanming Wang, Ruifeng Liang, Youcai Tuo, Kefeng Li, and Ben Hodges. 77:478-484.

Impact of Minimum Daily Dissolved Oxygen Concentration on Production Performance of Hybrid Female Channel Catfish × Male Blue Catfish. Les Torrans, Brian Ott, and Brian Bosworth. 77:485-490.

[Communication] Density-Dependent Impacts on Growth and Body Condition of Red Drum in Stock Enhancement Rearing Ponds. Joel D. Anderson and Paul D. Cason. 77:491- 496.

[Communication] Lower Lethal Temperature for Arapaima Arapaima gigas: Potential Implications for Culture and Establishment in Florida. Larry L. Lawson, Quenton M. Tuckett, Katelyn M. Lawson, Craig A. Watson, and Jeffrey E. Hill. 77:497-502.

566 Fisheries | Vol. 40 • No. 11 • November 2015 MIGRATION ECOLOGY It’s 3:00 a.m. OF MARINE FISHES David Hallock Secor “Tour de force synthesis on marine Do you know fish migrations. An essential read for ecologists of all specialties. “ —Michael Sinclair, author of Marine where your Populations: An Essay on Population Regulation and Speciation fish are? $99.95 hardcover / ebook

FRESHWATER FISHES OF NORTH AMERICA Volume 1: Petromyzontidae With technical expertise to Catostomidae that spans nearly all facets edited by Melvin L. Warren, of fisheries telemetry, we Jr., and Brooks M. Burr illustrated by Joseph R. Tomelleri are happy to share what “The first volume of a highly anticipat- we’ve learned. Contact us ed three-volume set that may be the for a free consultation to most important fish books on North discuss your project and American fishes produced in the last decade.”—Copeia your needs. $100.00 hardcover / ebook Downloaded by [Department Of Fisheries] at 23:47 13 March 2016

1-800-537-5487 / press.jhu.edu blueleafenviro.com

Fisheries | www.fisheries.org 567 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 CALENDAR will also be printed in Fisheries magazine.) More events listed at www.fisheries.org

January 24–27, 2016 76th Midwest Fish & Wildlife Conference | Grand Rapids, Michigan | midwestfw.org

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

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

May 21, 2016 2nd World Fish Migration Day | www.worldfishigrationday.com

May 23–27, 2016 Piscicide Course | USU, Logan, Utah

May 23–27, 2016 7th World Fisheries Congress | Busan, South Korea | wfc2016.or.kr

June 12–16, 2016 12th International Congress on the Biology of Fish | San Marcos, Texas | txstate.edu/continuinged/Events/ICBF.html

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

CORRECTION

Errata for Fisheries article, “Conserving Pacific Lamprey through Collaborative Efforts” Volume 40, Issue 2, February 2015, pages 72-79. doi: 10.1080/03632415.2014.996871

The below corrects a few errors noted in the original publication.

1. Abstract – change sentence, “To assess and conserve Pacific Lamprey, we …” to “To assess and conserve Pacific Lamprey, the Conservation Initiative partnership…” 2. Page 73 – last paragraph, change sentence, “In order to inform a comprehensive conservation plan, our first step…” to “In Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 order to inform a comprehensive conservation plan, the partnership’s first step…” 3. Page 74 – Conservation Initiative section, first paragraph, add (Goodman and Reid 2012) to sentence, “We applied the Strategic Habitat Conservation (SHC) approach …for Conservation Measures (Assessment; Luzier et al. 2011, Goodman and Reid 2012)…” 4. Page 75 – First paragraph, change sentence, “The RMUs are…” to “The RMUs were…and Alaska (Figure 1), since the signing of the Conservation Agreement, California has further subdivided the RMUs into seven.” 5. Page 76 – Implementation Plans paragraph, change sentence from, “We are in the process of developing…for each of the 10 RMUs.” to “We are in the process of developing…for each of the 15 RMUs.” 6. Page 76 – Results section, second paragraph, change sentence starting, “In Northern California…” change to “According to Goodman and Reid 2012, north of Point Conception Pacific Lamprey were…” 7. Figure 2 caption – Add (Goodman and Reid 2012). Change NA to read, “NA is for HUCs with insufficient data and for river basins in California with no access to the ocean.” 8. Acknowledgements – change first sentence to read, “We thank the members of the USFWS Western Lampreys Conservation Team with whom we work jointly on the Initiative.”

568 Fisheries | Vol. 40 • No. 11 • November 2015 Dear Members,

Thank you for your support and participation. We could not have reached our 2015 goals without you.

The 2016 renewal period is now open. Keeping your membership current is an easy online process—just log into your AFS account and click "renew." Please renew your membership dues before the end of this year to ensure no interruptions in your benefits, including receipt of our bi- weekly newsletter, online access to Fisheries and our directories, and bookstore discounts.

Show your support for the world’s oldest society dedicated to fisheries science with your prompt renewal! We look forward to another successful year in 2016.

Thanks again! Your Membership Team—Eva and Juanita Downloaded by [Department Of Fisheries] at 23:47 13 March 2016

Fisheries | www.fisheries.org 569 TAXONOMIC SERVICES WWW.ECOANALYSTS.COM The World Leader & Innovator in Fish Tags 20 YEARS In honor of our 20th anniversary, EcoAnalysts is pleased to announce that we are offering a significant discount (up to 50%) on taxonomy services. (Based on a 90 day Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 processing �me with data delivery falling between April 1 - July 30, 2016.) Available capacity is limited first come, first served. Call or email for EcoAnalysts premium taxonomy services. • Call 800-843-1172 to discuss your custom tagging needs • Email us at [email protected]

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570 Fisheries | Vol. 40 • No. 11 • November 2015 floy tag ad3.indd 1 1/24/2013 6:45:34 PM BACK PAGE

The smile of the Sheepshead. Photo credit: Joshua Raabe. The Teeth of Fishes: Say Ahhh! Natalie Sopinka AFS Contributing Writer E-mail: [email protected]

A dentist would be quite impressed with the array of tooth Dentists may be perplexed when it comes to American types among fishes. Paddlefish Polyodon spathula. The genus Polydon translates For coral-crunching Queen ParrotfishScarus vetula, the to “many teeth” in Greek, yet adults have no teeth. American upper and lower teeth are fused together and to their jawbones, Paddlefish do develop teeth as larvae when genes associated forming two sturdy plates that resemble a parrot’s beak. with tooth development are turned on (Smith et al. 2015). As Carnivorous fishes have treacherous canine teeth. The Great larvae grow older, tooth development slows and teeth are lost. Barracuda Sphyraena barracuda has prominent, vampire-like So what does the genus name refer to? Consensus is that the fangs. Looking for cavities in the mouth of the aptly named “many teeth” are the many gill rakers that adult paddlefish make Toothed Seadevil Neoceratias spinifer, a species of anglerfish, use of when filter-feeding. would be a terrifying ordeal. Their mouths are lined with Finally, for those dentists that are also history buffs, jaggedly arranged needlelike (or villiform) teeth effective the fossil fish Romundina stellina is sure to make a lasting at caging prey in their mouths. Round Goby Neogobius impression. Around 400 million years ago, R. stellina prowled melanostomus have a relatively harmless grin. Their hidden the Earth’s waters with teeth made of materials similar to those molariform teeth located in their throats (known as pharyngeal found in our human teeth (e.g., an enamel-like protective cap; teeth) are what makes them zebra mussel-crushing aficionados Rücklin and Donoghue 2015). If human teeth evolved from (Ghedotti et al. 1995). Flossing out benthic invertebrates must fish teeth, what came before fish teeth? The leading hypothesis Downloaded by [Department Of Fisheries] at 23:47 13 March 2016 be rather tedious for Blue CatfishIctalurus furcatus. Their is that the teeth of fish evolved from the scales of fish (the repeating rows of tiny (or cardiform) teeth resemble the wire “outside-in” hypothesis; Donoghue and Rücklin 2014). brushes we use to groom cats and dogs. Common Pike Eel Wouldn’t you know? Fish were the first animals that would Muraenesox bagio have triangular teeth on a bone that protrudes have needed to visit the dentist’s office. from the roof of their mouth (known as vomerine teeth). A dentist would be shocked when peering into the mouth REFERENCES of a Sheepshead Archosargus probatocephalus or Black Pacu Correia, J. P. 1999. Tooth loss rate from two captive Sandtiger Sharks Colossoma macropomum. Their teeth are freakishly human-like: (Carcharias taurus). Zoo Biology 18(4):313-317. Donoghue, P. C. J., and M. Rücklin. 2014. The ins and outs of the evo- incisors in the front, molars in the back. Sheepshead’s teeth are lutionary origin of teeth. Evolution and Development. DOI:10.1111/ used to shatter shelled invertebrates, whereas Black Pacu chomp ede.12099. away on the seeds of tropical plants. Ghedotti, M. J., J. C. Smihula, and G. R. Smith. 1995. Zebra mussel The rates of tooth loss (~1 per day) in Sand Tiger Sharks predation by Round Gobies in the laboratory. Journal of Great Lakes Research 21(4):665-669. Carcharias taurus (Correia 1999) might alarm a dentist at first. Rücklin, M., and P. C. Donoghue. 2015. Romundina and the evolution- But sharks are always growing new teeth to replace old teeth. ary origin of teeth. Biology Letters 11(6):20150326. The rotating rows of teeth means these polyphyodont predators Smith, M. M., Z. Johanson, T. Butts, R. Ericsson, M. Modrell, F. J. Tu- lenko, M. C. Davis, and G. J. Fraser. 2015. Making teeth to order: are always equipped for capturing a meal. conserved genes reveal an ancient molecular pattern in paddlefish (Actinopterygii). Proceedings of the Royal Society of Lon- don B: Biological Sciences 282(1805):20142700.

Fisheries | www.fisheries.org 571 Our transmitters aren’t as interesting as what researchers put them on.

Golden Mahseer in Bhutan. Photo credit: Karl Anderson. Teeth of the Sand Tiger Shark. Photo Credit: Amanda Pippin. Barracuda - say cheese! Photo Credit: Florida Fish and Wildlife Con- servation Commission.

A smiling Queen Parrotfish. Photo Credit: Kevin Bryant. The smile of the Round Goby. Photo credit: Julie Marentette.

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572 Fisheries | Vol. 40 • No. 11 • November 2015 Our transmitters aren’t as interesting as what researchers put them on.

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Kelp forest image courtesy Kevin Panizza