ASIH PRESIDENTIAL ADDRESS Copeia 105, No. 1, 2017, 10–13
GIANTS! Or. . .The Return of the Kelp Forest King Larry G. Allen1
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NE typically starts such a plenary address by waxing philosophical. This address will be no exception, but I will be brief in this endeavor. It has been a great honor and pleasure to have served as President of our 100year-old society. I became a member of ASIH in 1972 as an undergraduate at California State University Fullerton. To this day, ASIH remains the society that I care about the most. Back in 1972 it never occurred to me that I would, one day, have the privilege of serving ASIH as President. To add my name to the list of past presidents, Giants of our field like Carl Hubbs, Robert Miller, Boyd Walker, Bruce Collette, Jay Savage, Dan Cohen, Clark Hubbs, Dick Rosenblatt, Ted Pietsch, and Bob Cashner, to name a just a few, was truly a dream come true. Philosophically, I am a naturalist and a teacher. As such, I use whatever tools I can find to answer the questions that I find interesting about fishes. When I became a faculty member at what was at the time, a ‘‘teaching institution’’ in 1982, I realized that my students were the key to answering these questions. To this day, despite institutional road blocks, I steadfastly hold that field research is the best teaching tool we have at our disposal. I also believe that plenary presentations, like lectures, should be short, informative, and entertaining. So, for this plenary talk, I chose to tell the story of my lab’s recent studies on another kind of Giant. In producing this talk, I liberally spaced underwater video of Giant Sea Bass throughout to allow the audience to ‘‘be one with the fishes’’ (my credo to all of my students). Unfortunately, video does not translate well into the present article format, but I will carry on nonetheless. We begin our story about Giant Sea Bass over 125 years ago in the 1890s. Charles Frederick Holder, a prominent citizen in Southern California about the turn of the 20th century, moved from the American Natural History Museum in New York, and took up residence in Pasadena. In California he became a successful business man and, among other things, helped establish the Rose Parade. He was also an avid angler and started writing popular articles about angling and the outdoors of California to help attract tourists. In an early Scribner’s Magazine article he wrote about Giant Sea Bass off Catalina. Black sea bass is the name that Holder used and that is still the name that the recreational anglers use. The proper common name changed to Giant Sea Bass in 1960 to avoid confusion with the black sea bass (Centropristis striatus) from the coastal regions of the eastern United States. The Haunt of the Black Sea Bass It must be taken free-handed, a fight at arm’s-length, and being such, the moments fly by; it is half an hour, and we 1
have not yet seen the outline of our game. . . ‘‘It take your wind,’’ said Joe, with a low laugh. So it had, and I stood braced against the gunwale after a final dash—a burst of speed—to see a magnificent fish, black. . . pass swiftly across the line of vision, whirling the boat around end for end. ‘‘You’ve got him,’’ from astern, is encouraging; yet I have my doubts;. . . I was in the toils. But the flurry was the last. Several sweeps around the boat, and the black sea-bass lay alongside, covering boat and men with flying spray with strokes of its powerful tail. ‘‘It is a small one,’’ ejaculates my man, wiping the spray from his face. —Charles Frederick Holder, Scribner’s Magazine, 1891 The passage above represents how Holder was writing about this fish way back then. Anglers were catching these huge fish on hand lines. I love the liquid prose therein. The description that the fish takes his wind and that he is in his toils are simply classic. The fish finally approaches and starts turning the boat around end to end. Its powerful tail starts splashing water all over them. Then his pangero Joe says simply it’s a small one. I love all that. This story took place off Avalon at Catalina Island. So recreational anglers were catching these regularly at Catalina off of Avalon over one hundred years ago (Fig. 1). It just so happens that it is not far from where we study them now. In his 1910 work, A Book for the Angler, Sportsman and Tourist of California, Holder further characterized the Giant Sea Bass thusly, ‘‘The black sea bass when alive is a handsome fish of the black-bass type, fin for fin. In the water he looks blue, but is, in reality, a dark mahogany tint above, lighter beneath. The eye is large, and blue. All in all, the fish is a dignified and attractive fellow.’’ I agree. Holder also established the famous Avalon Tuna Club in 1898. So, at one time the Giant Sea Bass, along with world class Bluefin Tuna, attracted fisherman from all over the world to Southern California. And they came here simply to catch these big fish, particularly these trophy-sized bluefin. One of Holder’s very good friends and resident of Avalon for a time, Zane Grey, also wrote many articles on fishing of bluefin and black sea bass off of Catalina in those days as well (Grey, 1919, 1952). So, there is some historical perspective. Now, what about the fish itself? Giant Sea Bass, Stereolepis gigas (Fig. 2), is a member of the wreckfish family, Polyprionidae, based on larval characteristics and genetic analysis. The Polyprionidae includes two genera (Polyprion and Stereolepis) with two species of Stereolepis, one off of California and the other off of Japan. These
Department of Biology, California State University Northridge, Northridge, California; Email:
[email protected]. DOI: 10.1643/CI-17-577 Published online: 10 March 2017 Ó 2017 by the American Society of Ichthyologists and Herpetologists
Allen—ASIH Presidential Address
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Fig. 3. Photograph of a YOY Giant Sea Bass (7.2 cm) about five months old (photo credit Kevin Lewand, Monterey Bay Aquarium).
Fig. 1. A (then) world record (428 pound) Giant Sea Bass (Stereolepis gigas) caught by John T. Perkins off Catalina Island, California in June 1905 in the years prior to the fishery collapse for this apex predator off California in 1934.
fish get really big, obviously; we wouldn’t call them Giants if they weren’t giant. They have, up until our work recently, been reported as being up to 2.1 meters total length, and 255 kilograms. For those fishermen among us, that’s seven feet
Fig. 2. The largest Giant Sea Bass ever measured (2.7 m TL). Photograph taken at Goat Harbor, Santa Catalina Island in June 2015 by Parker House with J. R. Clark in background for scale.
and 560 pounds. Giants start out life on the bottom looking very different from adults (Fig. 3). As a matter of fact, all wreckfishes have very different juvenile forms. Giant Sea Bass, in the Northeast Pacific, occur from about Humboldt Bay south to all the way down around Baja, and up into the northern Gulf of California. However, on the Pacific coast, Giants rarely occur north of Point Conception and function as apex predators of the kelp bed and nearshore rocky reefs off the Californias (Horn and Ferry, 2006)—an apex predator that has been missing from the kelp forest and deep rock reefs of California’s nearshore ecosystem for almost a hundred years. So what happened to the Giants? At the end of the 1920s, the commercial fishing fleet out of San Pedro, California was growing and began fishing (gill netting and hand lining) Giant Sea Bass in earnest, often targeting their spawning aggregations in the summer months (Fig. 4). This practice led to nearly complete demise of the Giant Sea Bass fishery off California by 1934 (Croker, 1937). Since 1934, the only appreciable commercial landings of Giant Sea Bass were from off Baja California. Their commercial landings and populations decreased to almost nothing until, in 1982, the Mexican government prohibited the commercial take of Giants from Mexican waters. In the same year, the California Department of Fish and Game placed a complete moratorium on the recreational catch, and commercial catch was limited to two fish and later to one per trip. Then, in 1990, an epic thing happened in California. In a win for conservation, the voters of California passed Proposition 132, which banned all gill netting from in-shore waters and up to three miles off shore. Its implementation in 1994 changed everything (Pondella and Allen, 2008). How about now? Recreational take of Giants has not been allowed since 1982, but commercial fishers can still take one fish per trip. In the bygone days, a trip offshore or off Baja may have lasted two weeks. Now a trip typically lasts one day. So commercial fisherman can still take these fish. I admit to being torn because this incidental commercial take provided many of the specimens we studied in our lab, but these regulations still allow the legal take of an IUCN critically endangered species. One of the reasons that they remain vulnerable is that they aggregate to spawn. Over the last two decades, there have been indications that Giants were making a comeback. Various knowledgeable observers started to see more and more of them at historical spawning
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Copeia 105, No. 1, 2017
Fig. 4. Commercial landings of Giant Sea Bass from California waters from 1928 to 2008 (source: California Department of Fish and Wildlife).
aggregations sites. The number of underwater photographs of recent aggregations available online skyrocketed. Many talented underwater photographers began to share their photos of Giants at places like La Jolla, Catalina Island, Palos Verdes and Anacapa Island within the southern California Bight. So, we had anecdotal evidence that the historical spawning aggregations were returning. At that point, we began looking for other lines of evidence indicating a recovery was underway. One strong line of evidence came from a 15-year gill netting study targeting White Seabass that my group was conducting with Dan Pondella’s group at Occidental College, California. White Seabass are very large croakers that were similarly overfished and a hatchery enhancement program was launched in 1985. That program, the Ocean Resources Enhancement and Hatchery Program (OREHP), contracted us to go out and collect the tagged fish released from the hatchery using set gill nets throughout the Southern California Bight. The goal was to catch juvenile White Seabass, and the gill net protocol did a very good job collecting nearly 11,000 juvenile White Seabass. During the 15 years of this sampling program, we also started to catch juvenile Giant Sea Bass in these small, variable mesh gill nets. Ultimately, we found a significant increase in catch of juvenile Giant Sea Bass (Pondella and Allen, 2008) over the 15-year period. Additionally, two time series of established SCUBA surveys off of Palos Verdes, California conducted by Occidental College and off of Catalina Island, near Avalon (House et al., 2016) dating back into the early 1970s both showed a clear appearance of Giants suddenly between 2000 and 2003. From these three fishery-independent sources of information, I feel comfortable in stating that, yes, indeed, the Giants are returning. So, my message here is, if you protect them, they will return. What specifically brought them back? The gill net ban in 1994 allowed the comeback of these magnificent creatures, as well as the White Seabass and three species of sharks (soupfin, leopard, and gray smoothhound; Pondella and Allen, 2008). Then in January 2010, something serendipitous occurred. One of my commercial gill netter contacts, Gary Burke, called me. He said, ‘‘They have a big fish, a 500-pound fish, up at the docks in Santa Barbara. Do you want it?’’ I said no, but I
definitely wanted the skull! So, I drove to Santa Barbara that afternoon and purchased the head for $50. I then placed this 0.6 m head in the trunk of my car and headed home. From that point on, I continued to buy heads from the Santa Barbara Fish Market as well as the San Pedro Fish Market. These heads were the foundation for our age and growth studies on Giant Sea Bass. Power tools were necessary to extract the sagittal otoliths from these large, amazing skulls. It’s like getting into the skull of a mastodon or something like it. We eventually accumulated otoliths from 64 specimens that we sectioned and counted rings on digital images and ultimately aged fish from one to 76 years of age (Hawk and Allen, 2014). The first large fish from January 2010 was aged at 62 years. We were actually able to validate that the rings of this fish were indeed annual rings by analyzing the bomb radiocarbon at the focus and comparing that with the northeast Pacific bomb radiocarbon calibration chart. The bomb radiocarbon level matched 1949 which was the predicted year of birth of the 62-year-old specimen (Allen and Andrews, 2012). Using all of the age and length data (adjusting from head length to standard length) available, my student Holly Hawk and I were able to calculate the first the first von Bertalanffy growth curve (Hawk and Allen, 2014) on this species—ever. Next, because we had heads, we also had tissue archived for genetic analysis. Ultimately, we had 61 samples covering the primary range of the Giant Sea Bass from southern and Baja California, Mexico. Working with Holly Hawk and my postdoc, Chris Chabot, we carried out standard population genetics assays including the mitochondrial control region and microsatellite analysis of nuclear DNA (Chabot et al., 2015). The findings were surprising. We found that out of the 61 fish, we had only four mtDNA haplotypes. Anyway you cut it, this is very low genetic variability. All of the fish appeared to be close relatives. We also genotyped the 12 nuclear microsatellite loci. Again, our specimens represented a single genetic population regardless of whether they were taken from southern California or either side of Baja California. They also have a very low effective population size. It would only take 150 females to maintain this low
Allen—ASIH Presidential Address
genetic diversity. Over all we were forced to conclude that the Giants alive today represent a single genetically bottlenecked population. The source of the bottleneck? It is easy to conclude that the overfishing before the 1934 collapse had a lot to do with it. If so, then it has taken 90 years for the Giant Sea Bass population to recover to its present level, yet they are bottlenecked with low genetic diversity. Obviously this is not good for their long-term viability. So, where did we go from there? Our lab decided to begin studies of the aggregations occurring off of Catalina themselves. Because I was a USC alum, I have had strong connections with the Wrigley Marine Science Center for many years, and we had been sending summer fellowship students out there for a long time. So, I sent two of my students, Parker House and J. R. Clark, out to study giant sea bass in their natural habitat over the summers of 2014 and 2015. What did they find? Among other things, Giant Sea Bass regularly occur in pairs during the summer months. Parker and J. R. were out almost every day counting Giants, and 50% of the time they encountered them in pairs. Typically, one of the fish would be fairly rotund with a big white spot mid-flank above the vent. We believe these are females in spawning condition. The other member of the pair very often was positioned behind the rotund fish usually had a very sleek, almost concave abdomen. We think those are males. Parker and J. R. also started systematic sampling, running 5,000 square-meter stations around Catalina Island and using GoPro cameras mounted with parallel laser system with ten centimeters apart for the purpose of measuring individuals. Divers counted and measured Giants on 5, 100meter transects looking ten meters on both sides at eight stations around Catalina in 2014 and 2015. Where were the Giants off Catalina Island? It turned out that Italian Gardens and Goat Harbor were the sites where Giant Sea Bass were consistently abundant. This area just happens to now be a State Marine Reserve (a marine protected area) in California. It now appears that this ‘‘Long Point Marine Reserve’’ was established in the correct area to protect these aggregated Giant Sea Bass. Also, density estimates indicated that there were only about 40 to 50 individual Giants present off Catalina during those two summers (House et al., 2016). With length data derived from parallel lasers, we also were able calculate the first length-frequency analyses on Giants— ever. All of the measurements taken together did create duplication of individuals from month to month, so the data must be evaluated with caution. However, some general findings did emerge. We think that virtually all the fish encountered were mature individuals based on their lengths (.1 m). The second interesting outcome was that one very large fish, measured in both 2014 and 2015 and recognized as the same individual by spot and scar recognition markings, turned out to be the largest Giant Sea Bass ever measured. This humongous fish (Fig. 2) was estimated to be about nine feet long. This observation matches some old musings of Charles Holder and that some of the others have written about. The final general outcome from being able to measure lengths accurately was that we were able to estimate age from total lengths using Hawk and Allen (2014). From age we could back-calculate the year of birth and estimate
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year-class strength. We concluded that the majority (60%) of the Giants seen at Catalina Island had recruited after the gill net ban of 20 years ago. It seems that juvenile Giants are now released from the high mortality as bycatch in the old nearshore gill net fishery. What comes next? Two of my current graduate students both presented papers at the NOLA 100th Anniversary Meetings. J. R. Clark presented his findings on the courtship behavior of Giants, and Stephanie Benseman gave her talk on the distribution and abundance of young-of-year, ‘‘baby’’ Giants at these meetings. My research program has also begun a collaboration with Chris Lowe and his students of the CSULB SharkLab to pursue acoustic tagging studies of Giants throughout their range. So far, in 2015, we were able to place acoustic tags on six of them we have studied at Goat Harbor, Catalina Island. Preliminary results indicate that they move around a lot even when they are found in aggregations. Finally, as is evident throughout this presentation, most of my research contributions over the last 35 years would not have been possible without my students. I owe it all to them. Thank you, guys. In closing, I believe ASIH also owes a great debt to our student members, and I have tried everything in my power during my time as ASIH President to make attendance more affordable for the students. I think we are well on our way to accomplishing that goal. I feel that I am ‘‘preaching to the choir’’ when I reiterate that the students are our future! LITERATURE CITED Allen, L. G., and A. H. Andrews. 2012. Bomb radiocarbon dating and estimated longevity of Giant Sea Bass (Stereolepis gigas). Bulletin of the Southern California Academy of Sciences 111:1–14. Chabot, C. L., H. A. Hawk, and L. G. Allen. 2015. Low contemporary effective population size detected in the Critically Endangered Giant Sea Bass, Stereolepis gigas. Fisheries Research 172:71–78. Croker, R. S. 1937. Black sea bass. In: Bureau of Commercial Fisheries. The Commercial Fish Catch of California for the Year 1935. CFG Fish Bulletin 49:76–77. Grey, Z. 1919. Great Game Fishing at Catalina. Santa Catalina Island Co., Magill-Weinsheimer, Chicago. Grey, Z. 1952. Zane Grey’s Adventures in Fishing. Harper & Brothers, New York. Hawk, H. A., and L. G. Allen. 2014. Age and growth of the giant sea bass, Stereolepis gigas. CalCOFI Report 55:1–7. Horn, M., and L. Ferry-Graham. 2006. Feeding mechanisms and trophic interactions, p. 387–410. In: The Ecology of Marine Fishes: California and Adjacent Waters. L. G. Allen, D. J. Pondella, and M. H. Horn (eds.). University of California Press, Berkeley. House, P. H., B. L. F. Clark, and L. G. Allen. 2016. The return of the king of the kelp forest: distribution, abundance, and biomass of giant sea bass (Stereolepis gigas) off Santa Catalina Island, California, 2014–2015. Bulletin of the Southern California Academy of Sciences 115:1–14. Pondella, D. J., II, and L. G. Allen. 2008. The decline and recovery of four predatory fishes from the Southern California Bight. Marine Biology 154:307–313.
