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Using demographic analysis to assess the population size of shark species: a test using the White Shark (Carcharodon carcharias) sub-population off central.

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Presentation on theme: "Using demographic analysis to assess the population size of shark species: a test using the White Shark (Carcharodon carcharias) sub-population off central."— Presentation transcript:

1 Using demographic analysis to assess the population size of shark species: a test using the White Shark (Carcharodon carcharias) sub-population off central California, USA Gregor M. Cailliet 1, Kenneth J. Goldman 2,, Henry F. Mollet 1,3 1 Moss Landing Marine Laboratories, Moss Landing CA 95039-9647, USA 2 Alaska Department of Fish and Game, 3298 Douglas Place, Homer, AK 99603 USA 3 Monterey Bay Aquarium, 886 Cannery Row, Monterey, CA 93940-1023 USA Squillante in Ebert (2003) Acknowledgements: Karyn Brewster-Geisz, Steve Campana, Jason Cope, Enric Cortes, Dave Ebert, Malcolm Francis, Selina Heppell, Lynn McMassters, Chris Perle, Colin Simpfendorfer, Based upon:

2 White Sharks (Carcharodon carcharias) are: Difficult to sample because they are large & highly migratory (see below); Move both vertically and horizontally; and Tend to segregate by sex, age and size. Thus, their population or sub-population sizes are difficult to estimate. Jorgensen et al. (2010) and TOPP Tracking Data ( predators-topp) But some studies have tried to do this in several places around the world. This is our attempt. Ken Goldman Photo

3 A recent tag-recapture study (Chapple et al. 2011) used photographic identification of the trailing edge of the first dorsal fin as natural individual marks and used this to estimate the number of sub-adult and adult White Sharks presently in the central California sub-population. These authors used the results from a Bayesian hypergeometric model to conclude that the: Central California population consists of only 219 mature and sub-adult white sharks (highest- posterior density estimate); With a 95% credible interval of 130 and 275; Where sub-adults are defined as being sexually immature; and Older juveniles are found with adults in aggregation areas.

4 Chapple et al. (2011) further concluded that : This represented ~50% of the mature and sub-adult White Sharks for the entire eastern North Pacific Ocean (ENP); and When all life stages were considered, the total population of white sharks was “far lower” than that of other apex predators. This very low estimate created serious concern about the status of the California White Shark population, and consequently resulted in petitions to consider white sharks as a candidate for the U.S. (Federal Register / Vol. 77, No. 189 / Friday, September 28, 2012 / Proposed Rules) and State of California endangered species (ESA) lists. Under the Federal Endangered Species Act, an endangered species is defined as “any species which is in danger of extinction throughout all or a significant portion of its range.”

5 We wanted to re-assess this population estimate and demonstrate the utility of demographic analyses and age distribution as useful methods for estimating population sizes in sharks. Demography has been applied to several species of sharks, including White Sharks (Mollet & Cailliet 2002). For this purpose, one needs a reasonable idea of age structure, often gained from size structure, and some age-specific population parameters.

6 DEMOGRAPHIC ANALYSIS Was based upon an evaluation of existing and new life history information on the White Shark, Carcharodon carcharias: SIZE & AGE FREQUENCIES GROWTH RATES AGE-SPECIFIC REPRODUCTION MORTALITY ESTIMATES and STAGE-BASED MATRIX MODELS, AGE-BASED LESLIE MATRICES, and LIFE HISTORY TABLES (Mollet & Cailliet 2002)

7 DEMOGRAPHIC POPULATION MODEL ESTIMATES FOR ALL AGE CLASSES We used demographic parameters to assess the size of the White Shark (Carcharadon carcharias) population off central California because recent studies provided appropriate data but did not utilize a demographic approach. We aimed to estimate a new minimum total population estimate for White Sharks off California coast using demographics, life history matrix information, and a conservative extension of Chapple et al.’s (2011) estimate to include age classes that were either poorly sampled, or not sampled at all, by their methodology. We did not attempt estimate the total population of White Sharks throughout the ENP as there are insufficient length, age and abundance data on which to base demographic models for the entire region.

8 To re-assess Chapple et al.’s (2011) sub-population estimate to produce a population estimate for all life stages of White Sharks, we relied on well- established demographic methods based on previously published estimates of vital life history parameters (Mollet & Cailliet, 2002). Jorgensen et al. (2010) sampled 179 White Sharks from the three locations below (Tomales Point, SE Farallon Islands, and Año Nuevo Island). However Chapple et al. (2011) only used data from the first two locations. Therefore, for our analysis, we only used data from those locations, not Año Nuevo Island, most closely reflecting how their estimate of 219 mature and sub- adult sharks was derived. Ken Goldman Photo

9 AGE- & SIZE-BASED FREQUENCIES The size-frequency and sex of the sharks sampled by Chapple et al. (2011) can be inferred from Jorgensen et al. (2010) as both studies were run in parallel. We generated sex-specific size frequencies for the 130 White Sharks enumerated by Jorgensen et al. (2010), assuming that these are the same sharks sampled by the parallel study of Chapple et al. (2011), for which no data on individual sharks were provided. See the next slides for the data…

10 SUPPLEMENTARY MATERIAL Table S1. Electronic tags deployed and DNA sampled from white sharks (Carcharodon carcharias) in the North Eastern Pacific, 2000-2008 ­­­­­­­­­­­­­­­­­­___________________________________________________________________________________________ Shark#Tag dateLocation*TLSexPAT#ACU#HaplotypeOffshore Site † Pop-upTrack days ­­­­­­­­­­­­­­­­­­­­­­­­­­___________________________________________________________________________________________ 110/16/00FAR457M1H13-Apr-01179 210/16/00FAR518-2 310/16/00FAR488-3 410/16/00FAR518F4 511/09/00FAR381-5 611/09/00FAR518F6 711/05/01FAR457-7H/C3-May-02179 811/05/01FAR488F8C19-Jul-02256 911/05/01FAR427M9C14-Jul-02251 1011/05/01FAR442F109-Dec-0134 1111/15/01FAR450M11C11-Jun-02208 1211/15/01FAR380M1213-Jan-0259 1311/12/02FAR430M13 1411/15/02FAR530F14C14-Feb-0391 1509/24/03FAR427-15C22-Mar-04180 1610/08/03FAR442M16 1710/23/03FAR-F17 1810/27/03FAR396M18 1910/31/03FAR488-19C5-Jun-04218 2010/31/03FAR488M20 2110/13/04TOM381-21Scot 2211/05/04FAR-M22C6-Sep-05305 2311/05/04FAR457M23C4-Apr-05150 2411/05/04FAR457M24C7-Mar-05122 2511/05/04FAR427M25C10-May-05186 2611/15/04FAR--GU002302 2711/30/04FAR396F26H10-Jun-05192 2811/30/04FAR490F27C3-May-05154 2912/03/04FAR426F288-Feb-0567 3010/01/05ANO397-GU002303 3110/01/05ANO488-GU002303 3210/20/05ANO411-29C30-Apr-06192 3310/20/05ANO411M30 3411/02/05ANO411F31C29-Aug-06300 3511/05/05ANO488F32C2-Aug-06270 3611/06/05FAR488-33GU002304C29-May-06204 3711/09/05ANO411-34C7-Aug-06271 3811/10/05ANO--GU002303 3911/10/05FAR457M35C4-Aug-06267 4011/16/05ANO411F36H/C16-Oct-06334 4111/16/05ANO427-377-Jan-0652 4211/17/05FAR488F38GU002305C15-Aug-06271 Scot Anderson

11 4311/17/05FAR366M39 4411/18/05FAR396M42GU002307 4511/18/05ANO366M40GU002306C17-Jul-06241 4611/18/05ANO457F41GU002303C17-Sep-06303 4711/18/05ANO400-43C21-Jul-06245 4811/18/05ANO488-44C16-Sep-06302 4911/19/05FAR488M45GU002308C26-Aug-06280 5011/19/05FAR488F46C16-Sep-06301 5111/20/05ANO427-GU002309 5211/20/05ANO442-GU002307 5311/20/05FAR427M47C17-Aug-06270 5411/20/05FAR457M48C19-May-06180 5511/21/05FAR427M49GU002310C19-Aug-06271 5611/22/05FAR457M50C18-Jul-06238 5712/09/05TOM426-51 5812/15/05TOM472F52 5912/15/05ANO480F53H11-Oct-06300 6012/15/05ANO390M54C3-Aug-06231 6101/21/06ANO400F55H/C9-Nov-06292 6201/21/06ANO400F56H3-Sep-06225 6310/02/06FAR335M57GU00230719-Jan-07109 6410/03/06FAR488M59GU002307C12-Jun-07252 6510/03/06FAR518M58GU002303C27-Apr-07206 6610/05/06FAR-MGU002311 6710/09/06FAR366M63GU002314 6810/09/06FAR366M62GU002313H2-Mar-07144 6910/09/06FAR366M60GU002312 7010/09/06FAR488F61GU002303C5-Jun-07362 7110/10/06FAR488M65GU002316H7-Jan-0789 7210/10/06FAR510M64GU002315C6-Aug-07300 7310/10/06FAR427M67GU00230718-Dec-0669 7410/10/06FAR518M66GU002303C6-Aug-07300 7510/10/06FAR457M68GU002303 7610/10/06FAR427-69H20-Sep-07239 7710/10/06FAR427M709-Jan-0791 7810/11/06REY427M71 7910/14/06FAR518M1 8010/20/06FAR472M722GU002303 8110/23/06FAR488M3 8210/23/06FAR488M4 8310/23/06FAR427M5 8410/23/06FAR396M6GU002307 8510/31/06FAR--GU002307 8611/01/06FAR457-7GU002307H 8711/01/06FAR457-8GU002303 8811/01/06FAR533-739GU002307H/C29-Jul-07270 8911/01/06FAR488-10 9011/01/06FAR442F74C21-Jan-0781 9111/01/06FAR488F75C25-Jul-07266

12 9211/01/06FAR427F7613-Nov-0612 9311/05/06TOM457F7718-Nov-0613 9411/10/06ANO488F11 9511/10/06TOM441M7812GU002317C13-May-07164 9611/10/06ANO366M79C26-Mar-07136 9711/10/06ANO426F809-Jan-0760 9811/18/06ANO488F81 9911/20/06TOM503F13GU002307 10012/03/06ANO396F14 10112/03/06ANO473F15H 10212/03/06TOM396-82GU002307H/C18-Jul-07227 10312/05/06TOM488F16GU002307 10401/24/07ANO-F17GU002318 10501/24/07ANO427M18 10601/24/07TOM411-19H 10701/24/07TOM305-20GU002307 10801/24/07ANO457F83 10909/17/07FER*-MGU002321 11010/06/07ANO427M21GU002303 11110/06/07ANO427M8422GU002307 11210/07/07ANO427M85GU002319 11310/07/07FAR--GU002303 11410/07/07TOM400-86 11510/08/07TOM427F8723 11610/08/07FAR--GU002303 11710/11/07FAR427M24GU002303 11810/11/07FAR457M88GU002303 11910/13/07TOM300-25 12010/13/07FAR396M90GU002307 12110/13/07FAR457M89GU00230427-Nov-0745 12210/13/07FAR457M91C20-Dec-0768 12310/13/07FAR457M92C27-Jun-08258 12410/14/07ANO457F26 12510/14/07FAR457M932711-Dec-0758 12610/14/07ANO427-28 12710/19/07FAR488M29GU002307 12810/19/07FAR457M30GU002303 12910/19/07FAR427M94H18-Feb-08122 13010/22/07FAR477M31GU002303 13110/22/07FAR477M32GU002316 13210/22/07FAR411M33 13310/23/07FAR477M34H 13410/23/07FAR488M35 13510/23/07FAR-M36GU002303H 13610/24/07FAR518F95372-Dec-0739 13710/27/07FAR518F9638GU002320C12-Sep-08321 13810/27/07FAR518F39GU002303 13910/28/07TRI*457-40GU002307 14010/28/07FAR518-41

13 14111/01/07FAR--GU002303 14211/04/07TOM304F42 14311/05/07TOM366F43 14411/05/07TOM427-44 14511/12/07TOM366F45GU002307 14611/16/07TOM427F46 14711/24/07TOM351M47 148 11/25/07ANO472F48 14911/25/07ANO442- 97C2-Apr-08129 15012/14/07ANO457-49 15112/14/07ANO457-50GU002303 15201/17/08TOM335-51 15309/14/08TOM366F52 15409/14/08TOM442M53 15509/17/08TOM457M54 15609/24/08TOM366M55 15709/26/08TOM427M56 15809/27/08TOM442M57 15909/28/08TOM320F58 16010/01/08TOM259-59 16110/01/08TOM396F60 16210/06/08TOM396M61 16310/06/08TOM488M62 16410/12/08FAR-M63 16510/13/08FAR442M64 16610/13/08FAR396M65 16710/13/08FAR457M66 16810/17/08TOM488F67 16910/22/08REY396M68 17010/22/08REY457M69 17110/27/08FAR396M70 17210/29/08FAR427M71 17310/29/08FAR488M72 17410/30/08TOM427M73 17511/05/08TOM488F74 17611/06/08REY274M75 17711/07/08TOM427M76 17811/08/08TOM366F77 17911/12/08TOM396M78 *Location codes; SEFI = Southeast Farallon Island, TOM = Tomales Point, ANI = Año Nuevo Island, REY = Point Reyes, TRI = Trinidad Head, HUM = Humbolt Bay. † Offshore sites visited; C = Café, H = Hawaii.

14 The estimated lengths of the 130 sharks enumerated by Jorgensen et al. (2010) were binned at 30 cm (~1 ft) intervals. Lengths were converted to age using an established, although unvalidated, length-age relationship (Cailliet et al. 1985). We then used natural mortality and life history estimates based on an assumed longevity of 30 years and increased fertility to insure a stationary age distribution (Lambda = 1.0), based on life history information in Mollet & Cailliet (2002). This is similar to what Cailliet (1992) did for his demographic study of Leopard Sharks (Triakis semifasciata) off central California. Leopard Shark, Monterey Bay Aquarium

15 Theoretical White Shark Survivorship Curve & Stationary Age Composition Based upon Cailliet et al.’s (1985) VBGF and Mollet & Cailliet’s (2002) demographic analysis This allowed us to produce a survivorship curve and expected stationary age frequency distribution of White Sharks of all ages (life stages) for this sub-population. Age Estimate (years) Ken Goldman Note; Longevity estimates range from 30 to 60+ years

16 SIZE FREQUENCY DATA From Jorgensen et al. (2010) These size- and sex-frequency data, when compared to estimated sizes at maturity (males ~3.6 m, females at 4.5-5.0 m) reveal that they primarily sampled mature males plus a few sub-adults and small mature females. Their sex ratio was 78 males: 33 females (χ2 = 10.6176, p < 0.01). Large mature females were almost entirely absent from their sub-population estimate, and smaller size classes (pups, juveniles an sub-adults) were markedly under-represented. We then converted these sizes to ages so they could be plotted on a stationary age distribution curve like in the previous slide.

17 DEMOGRAPHY We used established demographic methodology (Mollet & Cailliet 2002) to generate a survivorship curve (stationary age distribution) using: An assumed longevity of 30 years, a very conservative estimate given existing growth functions; An instantaneous natural mortality estimate of M= 0.1535 (~85.7% survivorship) per year; Age at maturity = 15 years; and Annual fertility (number of female pups per year) = 1.553, a slightly higher rate than reported in the literature (e.g. Mollet & Cailliet 2002, Francis 1996). All to achieve a stationary age distribution (lambda = 1.0).

18 THE ABUNDANCE OF ALL LIFE STAGES We matched the stationary age distribution to the realized proportional age frequency of Jorgensen et al. (2010), scaled to reflect Chapple et al.’s (2011) estimate of 219; Then we matched the largest peaks in the sampled age frequency to the theoretical curve and applied the appropriate scaling factor to all other age classes, assuming that Chapple et al’s. (2011) estimate was correct for the most abundant age classes (12, 13, & 14); and Then, we applied the proportion of estimated white sharks at a given age to those of ALL OTHER AGE CLASSES, we were able to derive a numerical estimate for all age classes combined.

19 THE ABUNDANCE OF ALL LIFE STAGES (Continued) The “ALL OTHER AGE CLASSES” mentioned in the previous slide are represented in the grey shading below. (These are mainly pups, YOY, juveniles and large adults)

20 OUR RESULTS Chapple et al. (2011) only sampled a small proportion of the age distribution for the sub-population represented at the sampling locations. Matching the sampled age frequency of the most abundant age classes with neighboring age distribution proportions (at 12, 13 and 14 yrs) and scaling appropriately provides an estimate of number of all life stages (ages) throughout coastal California would be at least 2,418 White Sharks. The range was from 2,148 to 2,819 White Sharks, depending upon which size or age class we chose, with 2,418 as the final value with the best fit at age 13. Increasing M by a factor of two to reflect increased mortality of early life stages for each of the two youngest age classes results in a negligible effect (<2%) on the population estimate.

21 What if White Sharks Live Longer than 30 years? A recent study by Hamady et al. (2014) used bomb radiocarbon to validate the ages of 8 White Sharks from the western North Atlantic. Their results suggested that White Sharks could perhaps live up to 60, maybe 70 yrs (also theoretically predicted by Cailliet et al. (1985). This could then theoretically change the growth curves of White Sharks as follows: Squillante in Ebert (2003)

22 What if White Sharks Live Longer than 30 years (continued)? This would change the theoretical survivorship curve or stationary age distribution as seen below. However, it would only make our population estimate even larger (between ~3,450 to ~ 5,120 White Sharks; Burgess et al. 2014). Age Estimate (years)

23 Our new estimate is comparable to white shark population size estimates from other locations around the world (Burgess et al. 2014) * * * * = Discussed in later slides

24 In the eastern North Pacific (U.S. west coast): Our estimate is similar to that from a recent independent status review by the NMFS (Dewar et al. 2014) of the ENP white shark population, which estimated ~3,000 individuals. This status review team had access to virtually all available data from white shark researchers throughout the eastern North Pacific and used multiple modeling approaches. Similar conclusions were made in a 2014 analysis by the California Department of Fish and Wildlife (and approved by the Commission) Scot Anderson

25 In the western North Atlantic: Another study (Curtis et al. 2014) was just published indicating an increase in relative abundance of White Sharks: Seasonal distribution and historic trends in abundance of White Sharks, Carcharodon carcharias, in the western North Atlantic Ocean, by Curtis TH, McCandless CT, Carlson JK, Skomal GB, Kohler NE, et al. (2014) Seasonal Distribution and Historic Trends in Abundance of White Sharks, Carcharodon carcharias, in the Western North Atlantic Ocean. PLoS ONE 9(6): e99240. doi: 10.1371/journal.pone.0099240

26 In Australia: Malcolm et al. (2001) used catch and effort data, historical fishing records, shark control program records, cage-diving operation information, telemetry, tag and recapture, and shark attack data in a deterministic population size model to estimate a population size of between 2,728 and 13,746 female White Sharks. In another study, Blower et al.( 2012), used genetic techniques to estimate the effective population size of white sharks off southern Australia at approximately 1,512 individuals. This study also suggested that a minimum of 500-1,000 breeding individuals would be required to retain enough genetic variability and ensure evolutionary potential. Although the size of the central California coastline is significantly smaller than the area studies in Australia, the differences in population size estimates is notable. This also contrasts with statements by Chapple et al. (2011) that the size of the eastern North Pacific White Shark population is alarmingly low, and cited genetic diversity as a factor.

27 And, in South Africa: A mark-recapture study by Towner et al. (2013) using photo-ID techniques at two adjacent White Shark aggregation locations in Gaansbi (Gans Bay), South Africa resulted in an abundance estimate of 908 (808-1008). This study employed open-population models and used a common quantitative technique (qAIC) to compare between competing model parameterizations. This sub-population abundance is more than three times the Chapple et al. (2011) mark-recapture estimate, in a region quite biogeographically similar to Farallon Islands and Tomales Bay.

28 Accurately estimating total population sizes of sharks requires methodologies that account for biases introduced by sampling a limited number of sites and include all life history stages across the species’ range. Our results indicate that an all-life-stage sub-population size of at least 2,400 individual White Sharks in coastal California is required to account for the abundance of the 219 adults and sub-adults estimated in the original study. This is likely to be a conservative (under) estimate for the California sub-population. The true total White Shark population size throughout the eastern North Pacific is likely several times greater than any estimates as all studies exclude non-aggregating, or otherwise unobservable, sharks, and those that independently aggregate at other important eastern North Pacific sites. Other studies using a variety of methods indicate the same thing as our demographic approach. Our analysis indicates that White Shark abundance off central California is not in danger as previously proposed. Indications are that the ENP White Shark population is healthy and may be growing. CONCLUSIONS


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