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Reproductive Biology of Deep-Sea Catsharks (Chondrichthyes: Scyliorhinidae) in the Eastern North Pacific

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Reproductive Biology of Deep-Sea Catsharks (Chondrichthyes: Scyliorhinidae) in the Eastern North Pacific Environ Biol Fish (2008) 81:35–49 DOI 10.1007/s10641-006-9162-9 ORIGINAL PAPER Reproductive biology of deep-sea catsharks (Chondrichthyes: Scyliorhinidae) in the eastern North Pacific Brooke E. Flammang Æ David A. Ebert Æ Gregor M. Cailliet Received: 24 May 2006 / Accepted: 27 September 2006 / Published online: 14 November 2006 Ó Springer Science+Business Media B.V. 2006 Abstract Apristurus brunneus, A. kampae, and females reached TL50 at approximately 485 mm Parmaturus xaniurus were caught by trawl and TL and 490 mm TL, respectively. Parmaturus longline between Washington and southern xaniurus males reached TL50 at 444 mm TL and California (48°Nto32°N latitude) from June females at 501 mm TL. Gravid female A. brunn- 2001 through October 2004. In females, oviducal eus were found in all months of the year, but gland width increased with TL in all three species gravid P. xaniurus were found primarily June but remained within 4 to 6% of TL. Oocyte through September. Adult A. kampae females diameter increased with TL during reproductive were found only in July through November and development in all species, and continued to there were gravid females in all these months. increase with size after reproductive maturity was Based upon gonadosomatic indices, IG, these reached. The weight-length relationships differed species may mate within a defined period of time significantly among species but not between sexes (April through June); however, A. brunneus of any species from the eastern North Pacific appear to reproduce throughout the year and combined. Using logistic regression, it was deter- P. xaniurus reproduce within a defined season mined that total length at 50% maturity (TL50)of (approximately July through September). A. brunneus males was 514 mm TL and all A. brunneus males were mature at 547 mm TL. Keywords Maturity Á Seasonality Á Female A. brunneus reached TL50 at 501 mm TL Reproduction Á Egg case Á Weight–length and all female A. brunneus were mature at regression Á Gonadosomatic index 581 mm TL. Apristurus kampae males and B. E. Flammang Á D. A. Ebert Á G. M. Cailliet Introduction Pacific Shark Research Center, Moss Landing Marine Laboratories, Catsharks (Chondrichthyes: Scyliorhinidae) are 8272 Moss Landing Road, Moss Landing, the largest and the most diverse family of living CA 95039, USA sharks, totaling 16 genera and approximately 134 Present Address: species distributed worldwide (Compagno et al. B. E. Flammang (&) 2005). With the exception of morphological Department of Organismic and Evolutionary Biology, information provided in species descriptions, the Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA reproductive biology of the species within this e-mail: bfl[email protected] family is poorly known, despite their global 123 36 Environ Biol Fish (2008) 81:35–49 distribution (Springer 1979; Ebert et al. 2006). A major contributing factor to this is the depth at which many of these species are found (Springer 1979). In the eastern North Pacific (ENP), there are three deep-sea scyliorhinids reported in the deep-water slope habitats: brown catshark, Apri- sturus brunneus (Gilbert 1892), white-edge cat- shark, Apristurus kampae Taylor 1972, and filetail catshark, Parmaturus xaniurus (Gilbert 1892). The occurrence of these scyliorhinids as inci- dental catch in commercial fishing operations (Rogers and Ellis 2000; Ebert 2003) warrants research into their biology and distribution to gauge the need for fisheries management. There are no data on A. brunneus, A. kampae,or P. xaniurus between Cape Flattery, Washington (48°N) and Point Dume, California (34°N) for comparison with the studies conducted in more northern and southern regions (Jones and Geen 1977; Cross 1988; Balart et al. 2000). By examin- ing the biology of these deep-sea species through- out the extent of their range, we can better understand their life history. The aim of this project was to describe the reproductive biology of A. brunneus, A. kampae and P. xaniurus in the eastern North Pacific, between 48°N and 32°N latitude. Fig. 1 Catch locations of all Apristurus brunneus (n = 711), A. kampae (n = 97), and Parmaturus xaniurus Methods (n = 376) studied from the eastern North Pacific from June 2001 through October 2004. Inset is of catch locations of Field sampling trawl and longline surveys by NOAA Fisheries Northwest and Santa Cruz labs in Monterey Bay off central California from June 2001 through October 2004. Bathymetric Samples were obtained annually from fishery- contours are scaled at 500 m depths independent bottom trawl survey cruises con- ducted by the National Oceanic and Atmospheric Administration (NOAA) Fisheries Service three depth strata they defined as shelf (20– Northwest Fisheries Science Center (NWFSC) 183 m), shallow slope (184–549 m), and deep laboratories in Newport, OR and Seattle, WA. slope (550–1350 m), with the aim of sampling as The NWFSC provided samples from their annual many different habitat types as possible while slope and shelf Fisheries Research and Monitor- minimizing damage to trawl gear. ing (FRAM) cruises during the months of June The Federal Ecology Division (FED) of the through October from 2001 to 2004, between NOAA Fisheries Service Southwest Fisheries Cape Flattery, Washington (48°N latitude) and Science Center (SWFSC) laboratory in Santa San Diego, California (32°N latitude, Fig. 1). Six Cruz, CA provided samples obtained monthly by hundred sample locations were randomly chosen fishery-independent bottom trawl and longline each year from a map of the survey area divided survey cruises targeting commercial groundfish into grids 2 nautical miles by 1.5 nautical miles in species between Davenport, CA (approximately length. The NWFSC survey was designed to cover 37°N latitude) and Monterey, CA (approximately 123 Environ Biol Fish (2008) 81:35–49 37 36°N latitude) from June 2002 to March 2004 Sexual maturity of specimens was determined (Fig. 1, inset). Using a depth-stratified sampling by visual analysis of gonads and reproductive method, five stations at arbitrarily designated organs. In males, sexual maturity was determined depth gradients were surveyed monthly off by clasper development and calcification. Juvenile central California, dependent on weather. Bottom males had flaccid claspers that did not extend past trawls were conducted from 170 to 667 m depth. the posterior end of the pelvic fins. Adolescent Longlines had between 750 and 7250 hooks per males had claspers that had grown past the pelvic haul and longline haul depths ranged from 373 to fin edge but were not calcified. The terminal 503 m. No sampling was undertaken in the month clasper elements are fully calcified in mature of May of any year because SWFSC/FED individuals. Clasper length was measured to the resources were focused on another survey, the nearest mm along the inner length of the clasper, annual California Cooperative Oceanic Fisheries from the clasper origin at the apex of the cloaca, Investigations (CalCOFI) larval fish survey. to the distal tip of the clasper (Compagno 1984). Samples opportunistically collected by observ- Maturity status in females was determined by ers with the Pacific States Marine Fisheries vascularization of the ovary and development of Commission (PSMFC) between Monterey Bay the oviducal gland, which produces the egg case. (37°N) and Big Sur, California (36°N) were In juvenile females, there was no discernible dependent upon cooperation of individual com- oviducal gland and no oocytes were visible in the mercial fishing operations and observer availabil- ovary. Development of the small, heart-shaped ity. Additionally, specimens in collections at the oviducal gland began in adolescents. The oviducal California Academy of Sciences (CAS), Com- gland in adult females was large and bulbous, and monwealth Scientific and Industrial Research oocytes are ‡14 mm in diameter. The widths of Organization (CSIRO) Hobart, Tasmania, Los the uterus, the left oviducal gland, and mature Angeles County Museum (LACM), Museum of oocytes were measured to the nearest mm. In Comparative Zoology (MCZ), Scripps Institute of these scyliorhinids, only the right ovary was Oceanography (SIO), Smithsonian National developed and reproductively functional. The left Museum of Natural History (USNM), and the ovary was vestigial (Springer 1966). Ovary mass University of Washington (UW) were examined was recorded to the nearest tenth of a gram. to augment field samples. Institutional codes are Changes in gonad mass, mature oocyte diameter as designated in Leviton et al. (1985). and number, and liver mass were analyzed for developmental and seasonal variation. Biological data Egg case size and fecundity were evaluated for each species. Egg case length relative to female Field samples were frozen upon capture and TL was compared to determine if a relationship returned to Moss Landing Marine Laboratories exists, as purported in Cross (1988). The propor- (MLML) for examination. Specimens were tion of gravid females was calculated for each thawed, measured, and weighed. Total length month to investigate reproductive seasonality. (TL) was measured to the nearest millimeter Total length at first, 50% (TL50) and 100% (mm), with the tail of the specimen in line with (TL100) maturity was determined for males and the body (Ricker and Merriman 1945). Specimen females of all species. Size at first maturity was weight was determined to the nearest gram (g). determined to be the length of the smallest Geometric weight-length regressions were calcu- mature individual for each sex of each species. lated for males and females of each species using Size at 50% maturity was determined using the the equation W = aLb, where W is the weight in g, logistic equation detailed in Roa et al. (1999), À1 L is the total length in mm, and a and b are fitted PðLÞ¼að1 À eb0þb1LÞ , where P(L) is the pro- constants (Ricker 1979). Analysis of variance portion of mature individuals at length L, a is the (ANOVA) was used to determine difference in asymptote, b0 is the intercept, and b1 is the slope weight–length regressions among species, sex, and parameter.
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