Gear modifications for fishing octopus, Octopus vulgaris, on live-bottom and adjacent flat bottom habitats in coastal waters off North Carolina

The Common Octopus, Octopus vulgaris, is an r-selected mollusk found off the coast of North Carolina that interests commercial fishermen because of its market value and the cost-effectiveness of unbaited pots that can catch it. This study sought to: 1) determine those gear and environmental factors that influenced catch rates of octopi, and 2) evaluate the feasibility of small-scale commercial operations for this species. Pots were fished from August 2010 through September 2011 set in strings over hard and sandy bottom in waters from 18 to 30 m deep in Onslow Bay, N.C. Three pot types were fished in each string; octopus pots with- and without lids, and conch pots. Proportional catch was modeled as a function of gear design and environmental factors (location, soak time, bottom type, and sea surface water temperature) using binomially distributed generalized linear models (GLM’s); parsimony of each GLM was assessed with Akaike Information Criteria (AIC). A total of 229 octopi were caught throughout the study. Pots with lids, pots without lids, and conch pots caught an average of 0.15, 0.17, and 0.11 octopi, respectively, with high variability in catch rates for each pot type. The GLM that best fit the data described proportional catch as a function of sea surface temperature, soak time, and station; greatest proportional catches occurred over short soak times, warmest temperatures, and less well known reef areas. Due to operating expenses (fuel, crew time, and maintenance), low catch rates of octopi, and high gear loss, a directed fishery for this species is not economically feasible at the catch rates found in this study. The model fitting to determine factors most influential on catch rates should help fishermen determine seasons and gear soak times that are likely to maximize catch rates. Potting for octopi may be commercially practical as a supplemental activity when targeting demersal fish species that are found in similar habitats and depth ranges in coastal waters off North Carolina.

Historical knowledge of sharks: ancient science, earliest American encounters, and American science, fisheries, and utilization

In western civilization, the knowledge of the elasmobranch or selachian fishes (sharks and rays) begins with Aristotle (384–322 B.C.). Two of his extant works, the “Historia Animalium” and the “Generation of Animals,” both written about 330 B.C., demonstrate knowledge of elasmobranch fishes acquired by observation. Roman writers of works on natural history, such as Aelian and Pliny, who followed Aristotle, were compilers of available information. Their contribution was that they prevented the Greek knowledge from being lost, but they added few original observations. The fall of Rome, around 476 A.D., brought a period of economic regression and political chaos. These in turn brought intellectual thought to a standstill for nearly one thousand years, the period known as the Dark Ages. It would not be until the middle of the sixteenth century, well into the Renaissance, that knowledge of elasmobranchs would advance again. The works of Belon, Salviani, Rondelet, and Steno mark the beginnings of ichthyology, including the study of sharks and rays. The knowledge of sharks and rays increased slowly during and after the Renaissance, and the introduction of the Linnaean System of Nomenclature in 1735 marks the beginning of modern ichthyology. However, the first major work on sharks would not appear until the early nineteenth century. Knowledge acquired about sea animals usually follows their economic importance and exploitation, and this was also true with sharks. The first to learn about sharks in North America were the native fishermen who learned how, when, and where to catch them for food or for their oils. The early naturalists in America studied the land animals and plants; they had little interest in sharks. When faunistic works on fishes started to appear, naturalists just enumerated the species of sharks that they could discern. Throughout the U.S. colonial period, sharks were seldom utilized for food, although their liver oil or skins were often utilized. Throughout the nineteenth century, the Spiny Dogfish, Squalus acanthias, was the only shark species utilized in a large scale on both coasts. It was fished for its liver oil, which was used as a lubricant, and for lighting and tanning, and for its skin which was used as an abrasive. During the early part of the twentieth century, the Ocean Leather Company was started to process sea animals (primarily sharks) into leather, oil, fertilizer, fins, etc. The Ocean Leather Company enjoyed a monopoly on the shark leather industry for several decades. In 1937, the liver of the Soupfin Shark, Galeorhinus galeus, was found to be a rich source of vitamin A, and because the outbreak of World War II in 1938 interrupted the shipping of vitamin A from European sources, an intensive shark fishery soon developed along the U.S. West Coast. By 1939 the American shark leather fishery had transformed into the shark liver oil fishery of the early 1940’s, encompassing both coasts. By the late 1940’s, these fisheries were depleted because of overfishing and fishing in the nursery areas. Synthetic vitamin A appeared on the market in 1950, causing the fishery to be discontinued. During World War II, shark attacks on the survivors of sunken ships and downed aviators engendered the search for a shark repellent. This led to research aimed at understanding shark behavior and the sensory biology of sharks. From the late 1950’s to the 1980’s, funding from the Office of Naval Research was responsible for most of what was learned about the sensory biology of sharks.

Spatial and temporal variability in the distribution and abundance of larval and juvenile fishes associated with pelagic Sargassum

A survey of the larval and juvenile fishes associated with the pelagic Sargassum habitat in the South Atlantic Bight and adjacent western Atlantic Ocean was conducted from July 1991 through March 1993. Fishes representing 104 taxonomic categories were identified, including reef fishes, coastal demersal, coastal pelagic, epipelagic and mesopelagic species. The most important families were Balistidae and Carangidae, each represented by 15 species. Species composition, species diversity and abundance varied both seasonally and regionally. Diversity was highest during spring through fall over the outer continental shelf and in the Gulf Stream. Abundance decreased from spring through winter and from the continental shelf into offshore waters. The numbers of fishes and fish biomass were found to be positively correlated with the wet weight of algae in most cases examined. The results of this study will be useful to fisheries managers assessing the potential impacts of commercial Sargassum harvesting in the region.

A pilot survey of deepwater coral/sponge assemblages and their susceptibility to fishing/harvest impacts at the Olympic Coast National Marine Sanctuary (OCNMS). Cruise report for NOAA Ship McARTHUR II Cruise AR-04-04: Leg 2 (June 1-12, 2004)

The offshore shelf and canyon habitats of the OCNMS are areas of high primary productivity and biodiversity that support extensive groundfish fisheries. Recent acoustic surveys conducted in these waters have indicated the presence of hard-bottom substrates believed to harbor unique deep-sea coral and sponge assemblages. Such fauna are often associated with shallow tropical waters, however an increasing number of studies around the world have recorded them in deeper, cold-water habitats in both northern and southern latitudes. These habitats are of tremendous value as sites of recruitment for commercially important fishes. Yet, ironically, studies have shown how the gear used in offshore demersal fishing, as well as other commercial operations on the seafloor, can cause severe physical disturbances to resident benthic fauna. Due to their exposed structure, slow growth and recruitment rates, and long life spans, deep-sea corals and sponges may be especially vulnerable to such disturbances, requiring very long periods to recover. Potential effects of fishing and other commercial operations in such critical habitats, and the need to define appropriate strategies for the protection of these resources, have been identified as a high-priority management issue for the sanctuary.

Fish movement patterns in Virgin Islands National Park, Virgin Islands Coral Reef National Monument and adjacent waters

NOAA’s National Centers for Coastal Ocean Science (NCCOS)-Center for Coastal Monitoring and Assessment’s (CCMA) Biogeography Branch, National Park Service (NPS), US Geological Survey, and the University of Hawaii used acoustic telemetry to quantify spatial patterns and habitat affinities of reef fishes around the island of St. John, US Virgin Islands. The objective of the study was to define the movements of reef fishes among habitats within and between the Virgin Islands Coral Reef National Monument (VICRNM), the Virgin Islands National Park (VIIS), and Territorial waters surrounding St. John. In order to better understand species’ habitat utilization patterns among management regimes, we deployed an array of hydroacoustic receivers and acoustically tagged reef fishes. Thirty six receivers were deployed in shallow near-shore bays and across the shelf to depths of approximately 30 m. One hundred eighty four individual fishes were tagged representing 19 species from 10 different families with VEMCO V9-2L-R64K transmitters. The array provides fish movement information at fine (e.g., day-night and 100s meters within a bay) to broad spatial and temporal scales (multiple years and 1000s meters across the shelf). The long term multi-year tracking project provides direct evidence of connectivity across habitat types in the seascape and among management units. An important finding for management was that a number of individuals moved among management units (VICRNM, VINP, Territorial waters) and several snapper moved from near-shore protected areas to offshore shelf-edge spawning aggregations. However, most individuals spent the majority of their time with VIIS and VICRNM, with only a few wide-ranging species moving outside the management units. Five species of snappers (Lutjanidae) accounted for 31% of all individuals tagged, followed by three species of grunts (Haemulidae) accounting for an additional 23% of the total. No other family had more than a single species represented in the study. Bluestripe grunt (Haemulon sciurus) comprised 22% of all individuals tagged, followed by lane snappers (Lutjanus synagris) at 21%, bar jack (Carangoides ruber) at 11%, and saucereye porgy (Calamus calamus) at 10%. The largest individual tagged was a 70 cm TL nurse shark (Ginglymostoma cirratum), followed by a 65 cm mutton snapper (Lutjanus analis), a 47 cm bar jack, and a 41 cm dog snapper (Lutjanus jocu). The smallest individuals tagged were a 19 cm blue tang (Acanthurus coeruleus) and a 19.2 cm doctorfish (Acanthurus chirurgus). Of the 40 bluestriped grunt acoustically tagged, 73% were detected on the receiver array. The average days at large (DAL) was 249 (just over 8 months), with one individual detected for 930 days (over two and a half years). Lane snapper were the next most abundant species tagged (N = 38) with 89% detected on the array. The average days at large (DAL) was 221 with one individual detected for 351 days. Seventy-one percent of the bar jacks (N = 21) were detected on the array with the average DALs at 47 days. All of the mutton snapper (N = 12) were detected on the array with an average DAL of 273 and the longest at 784. The average maximum distance travelled (MDT) was ca. 2 km with large variations among species. Grunts, snappers, jacks, and porgies showed the greatest movements. Among all individuals across species, there was a positive and significant correlation between size of individuals and MDT and between DAL and MDT.

Cruise report: regional assessment of ecosystem condition and stressor impacts along the northwestern Gulf of Mexico Shelf. NOAA Ship 'Nancy Foster' NF-11-07-RACOW (August 8-16, 2011)

This cruise report is a summary of a field survey conducted along a portion of the U.S. continental shelf in northwestern Gulf of Mexico (GOM), at navigable depths along the coastline seaward to the shelf break (~100m) from about 89°30' W to 95°28' W longitude, August 8 – 16, 2011 on NOAA Ship Nancy Foster Cruise NF-11-07-RACOW. Synoptic sampling of multiple ecological indicators was conducted at each of 34 stations throughout these waters using a random probabilistic sampling design. The original study design consisted of 50 stations extending from the Mississippi delta all the way to the U.S./Mexican border, but vessel failures precluded sampling at 16 stations within the western-most portion of the study area. At each station samples were collected for the analysis of benthic community structure and composition; concentrations of chemical contaminants in sediments and target demersal biota; sediment toxicity; nutrient and chlorophyll levels in the water column; and other basic habitat characteristics such as salinity, temperature, dissolved oxygen, turbidity, pH, sediment grain size, and organic carbon content. Other indicators, from a human-dimension perspective, were also recorded, including presence of vessels, oil rigs, surface trash, visual oil sheens in sediments or water, marine mammals, or noxious/oily sediment odors. The overall purpose of the survey was to collect data to assess the status of ecosystem condition and potential stressor impacts throughout the region, based on these various indicators and corresponding management thresholds, and to provide this information as a baseline for determining how such conditions may be changing with time. While sample analysis is still ongoing, some preliminary results and observations are reported here. A final report will be completed once all data have been processed.

Cruise report: regional assessment of ecosystem condition and stressor impacts along the northeastern Gulf of Mexico shelf; NOAA Ship Nancy Foster NF-10-09-RACOW (August 13-21, 2010)

This cruise report is a summary of a field survey conducted along the continental shelf of the northeastern Gulf of Mexico (GOM), encompassing 70,062 square kilometers of productive marine habitats located between the Mississippi Delta and Tampa Bay, August 13–21, 2010 on NOAA Ship Nancy Foster Cruise NF-10-09-RACOW. Synoptic sampling of multiple ecological indicators was conducted at each of 50 stations throughout these waters using a random probabilistic sampling design. At each station samples were collected for the analysis of benthic community structure and composition; concentrations of chemical contaminants (metals, pesticides, TPHs, PAHs, PCBs, PBDEs) in sediments and target demersal biota; sediment toxicity; nutrient and chlorophyll levels in the water column; and other basic habitat characteristics such as depth, salinity, temperature, dissolved oxygen, turbidity, pH, CDOM fluorescence, sediment grain size, and organic carbon content. Discrete water samples were collected just below the sea surface, in addition to any deeper subsurface depths where there was an occurrence of suspicious CDOM fluorescence signals, and analyzed for total BTEX/TPH and carcinogenic PAHs using immunoassay test kits. Other indicators of potential value from a human-dimension perspective were also recorded, including presence of any vessels, oil rigs, surface trash, visual oil sheens in sediments or water, marine mammals, or noxious/oily sediment odors. The overall purpose of the survey was to collect data to assess the status of ecosystem condition and potential stressor impacts throughout the region, based on these various indicators and corresponding management thresholds, and to provide this information as a baseline for determining how such conditions may be changing with time. In addition to the original project goals, both the scientific scope and general location of this project are relevant to addressing potential ecological impacts of the Deepwater Horizon oil spill. While sample analysis is still ongoing, a few preliminary results and observations are reported here. A final report will be completed once all data have been processed.

Cruise report: summer 2009 Sapelo Island National Estuarine Research Reserve ecological characterization (June 7 - June 13, 2009)

This cruise report is a summary of a field survey conducted within the Sapelo Island National Estuarine Research Reserve (SINERR), located on the Georgia coastline, June 7 – June 13, 2009. Multiple indicators of ecological condition and human dimensions were sampled synoptically at each of 30 stations throughout SINERR using a random probabilistic sampling design. Samples were collected for the analysis of benthic community structure and composition; concentrations of chemical contaminants (metals, pesticides, PAHs, PCBs, PBDEs) in sediments and target demersal biota; nutrient and chlorophyll levels in the water column; bacterial contaminants in the water column; and other basic habitat characteristics such as depth, salinity, temperature, dissolved oxygen, turbidity, total suspended solids, pH, sediment grain size, and organic carbon content. In addition to the fish samples that were collected for analysis of chemical contaminants relative to human-health consumption limits, other human-dimension indicators were sampled as well including presence or absence of fishing gear, vessels, surface trash, and noxious sediment odors. The overall purpose of the survey was to collect data to assess the status of ecosystem condition and potential stressor impacts throughout SINERR, based on these various indicators and corresponding management thresholds, and to provide this information as a baseline for determining how such conditions may be changing with time. While sample analysis is still ongoing a few preliminary results and observations are reported here. A final report will be completed once all data have been processed. The results will provide a comprehensive weight-of-evidence basis for evaluating current condition (aka a “state-of-the-SINEER environmental report”) and serve as a quantitative benchmark for tracking any future changes due to either natural or human disturbances. Another goal of the study is to demonstrate its utility as a possible model for assessing the status of condition at other NEERS sites using similar and consistent methods to promote system-wide regional and national comparisons.

Fish lost at sea: the effect of soak time on pelagic longline catches

Our analyses of observer records reveal that abundance estimates are strongly influenced by the timing of longline operations in relation to dawn and dusk and soak time— the amount of time that baited hooks are available in the water. Catch data will underestimate the total mortality of several species because hooked animals are “lost at sea.” They fall off, are removed, or escape from the hook before the longline is retrieved. For example, longline segments with soak times of 20 hours were retrieved with fewer skipjack tuna and seabirds than segments with soak times of 5 hours. The mortality of some seabird species is up to 45% higher than previously estimated. The effects of soak time and timing vary considerably between species. Soak time and exposure to dusk periods have strong positive effects on the catch rates of many species. In particular, the catch rates of most shark and billfish species increase with soak time. At the end of longline retrieval, for example, expected catch rates for broadbill swordfish are four times those at the beginning of retrieval. Survival of the animal while it is hooked on the longline appears to be an important factor determining whether it is eventually brought on board the vessel. Catch rates of species that survive being hooked (e.g. blue shark) increase with soak time. In contrast, skipjack tuna and seabirds are usually dead at the time of retrieval. Their catch rates decline with time, perhaps because scavengers can easily remove hooked animals that are dead. The results of our study have important implications for fishery management and assessments that rely on longline catch data. A reduction in soak time since longlining commenced in the 1950s has introduced a systematic bias in estimates of mortality levels and abundance. The abundance of species like seabirds has been over-estimated in recent years. Simple modifications to procedures for data collection, such as recording the number of hooks retrieved without baits, would greatly improve mortality estimates.

Multidirectional movements of sportfish species between an estuarine no-take zone and surrounding waters of the Indian River Lagoon, Florida

We examined movement patterns of sportfish that were tagged in the northern Indian River Lagoon, Florida, between 1990 and 1999 to assess the degree of fish exchange between an estuarine no-take zone (NTZ) and surrounding waters. The tagged f ish were from seven species: red drum (Sciaenops ocellatus); black drum (Pogonias cromis); sheepshead (Archosargus probatocephalus); common snook (Centropomus undecimalis); spotted seatrout (Cynoscion nebulosus); bull shark (Carcharhinus leucas); and crevalle jack (Caranx hippos). A total of 403 tagged fish were recaptured during the study period, including 65 individuals that emigrated from the NTZ and 16 individuals that immigrated into the NTZ from surrounding waters of the lagoon. Migration distances between the original tagging location and the sites where emigrating fish were recaptured were from 0 to 150 km, and these migration distances appeared to be influenced by the proximity of the NTZ to spawning areas or other habitats that are important to specific life-history stages of individual species. Fish that immigrated into the NTZ moved distances ranging from approximately 10 to 75 km. Recapture rates for sportfish species that migrated across the NTZ boundary suggested that more individuals may move into the protected habitats than move out. These data demonstrated that although this estuarine no-take reserve can protect species from fishing, it may also serve to extract exploitable individuals from surrounding fisheries; therefore, if the no-take reserve does function to replenish surrounding fisheries, then increased egg production and larval export may be more important mechanisms of replenishment than the spillover of excess adults from the reserve into fishable areas.

Biology and population dynamics of cowcod (Sebastes levis) in the southern California Bight

Cowcod (Sebastes levis) is a large (100-cm-FL), long-lived (maximum observed age 55 yr) demersal rockfish taken in multispecies commercial and recreational fisheries off southern and central California. It lives at 20–500 m depth: adults (>44 cm TL) inhabit rocky areas at 90–300 m and juveniles inhabit fine sand and clay at 40–100 m. Both sexes have similar growth and maturity. Both sexes recruit to the fishery before reaching full maturity. Based on age and growth data, the natural mortality rate is about M =0.055/yr, but the estimate is uncertain. Biomass, recruitment, and mortality during 1951–98 were estimated in a delay-difference model with catch data and abundance indices. The same model gave less precise estimates for 1916–50 based on catch data and assumptions about virgin biomass and recruitment such as used in stock reduction analysis. Abundance indices, based on rare event data, included a habitat-area–weighted index of recreational catch per unit of fishing effort (CPUE index values were 0.003–0.07 fish per angler hour), a standardized index of proportion of positive tows in CalCOFI ichthyoplankton survey data (binomial errors, 0–13% positive tows/yr), and proportion of positive tows for juveniles in bottom trawl surveys (binomial errors, 0–30% positive tows/yr). Cowcod are overfished in the southern California Bight; biomass during the 1998 season was about 7% of the virgin level and recent catches have been near 20 metric tons (t)/yr. Projections based on recent recruitment levels indicate that biomass will decline at catch levels > 5 t/yr. Trend data indicate that recruitment will be poor in the near future. Recreational fishing effort in deep water has increased and has become more effective for catching cowcod. Areas with relatively high catch rates for cowcod are fewer and are farther offshore. Cowcod die after capture and cannot be released alive. Two areas recently closed to bottom fishing will help rebuild the cowcod stock.

The effect of intensive line fishing on the virgin biomass of a tropical deepwater snapper, the crimson jobfish (Pristipomoides filamentosus)

An intensive commercial hook-and-line fishing operation targeted the demersal fisheries resources at Saya de Malha Bank in the Southwest Indian Ocean. Fishing was conducted with 12 dories that were equipped with echo sounders and electric fishing reels and supported by a refrigerated mothership. Over a 13-day period in the 55–130 m depth range, a total of 74.3 metric tons (t) of fish were caught, of which the crimson jobfish (Pristipomoides filamentosus) represented 80%. Catch rates decreased with time and could not be attributed to changes in location, climatic conditions, fishing depth, fishing method, or bait type. The initial virgin biomass of P. filamentosus available to a line fishery at the North Western promontory of Saya de Malha Bank was estimated at 72.6 t through application of the Leslie model to daily catch and effort data. Biomass densities of 2364 kg/km2 and 1206 kg/km were obtained by applying the initial biomass estimates to the surface area and to the length of the dropoff that was fished. The potential sustainable yield prior to exploitation was estimated at 567 kg/km2 per year. The quantity of P. filamentosus caught by the mother-ship-dory fishing operation represented 82% of the initial biomass available to a hook-and-line fishery, equivalent to more that three times the estimated maximum sustainable yield. The results of the study are important to fisheries managers because they demonstrate that intensive line fishing operations have the potential to rapidly deplete demersal fisheries resources.

Migration patterns of spiny dogfish (Squalus acanthias) in the North Pacific Ocean

From 1978 to 1988, approximately 71,000 spiny dogfish (Squalus acanthias) were tagged off the west coast of Canada. This program is the most extensive tagging study conducted for a shark species. Twelve years after the last year of tagging, recaptured tagged spiny dogfish are still being reported. As of December 2000, 2940 tagged fish (4.1%) have been recaptured. Spiny dogfish were tagged in three major areas: Strait of Georgia, west coast Vancouver Island, and northern British Columbia waters. Generally, spiny dogfish were recaptured close to their release site; however, extensive migrations (up to 7000 km) did occur. Migration rates varied across release areas. Spiny dogfish tagged in the Strait of Georgia underwent the least extensive movement; only 10–14% of the recaptures occurred outside the strait. Spiny dogfish tagged off the west coast of Vancouver Island or in northern British Columbia waters underwent more extensive movement; approximately 49–80% of the tagged spiny dogfish recaptured outside of the release areas. Spiny dogfish from all three release areas were recaptured off the west coast of United States and Alaska. Most impressive are the recaptures of tagged spiny dogfish off the coast of Japan. Over 30 spiny dog-fish were recaptured near Japan, most of which originated off the west coast of Vancouver Island or from northern British Columbia waters.

The statistical properties of recreational catch rate data for some fish stocks off the northeast U.S. coast

Recreational fisheries in the waters off the northeast U.S. target a variety of pelagic and demersal fish species, and catch and effort data sampled from recreational fisheries are a critical component of the information used in resource evaluation and management. Standardized indices of stock abundance developed from recreational fishery catch rates are routinely used in stock assessments. The statistical properties of both simulated and empirical recreational fishery catch-rate data such as those collected by the National Marine Fisheries Service (NMFS) Marine Recreational Fishery Statistics Survey (MRFSS) are examined, and the potential effects of different assumptions about the error structure of the catch-rate frequency distributions in computing indices of stock abundance are evaluated. Recreational fishery catch distributions sampled by the MRFSS are highly contagious and overdispersed in relation to the normal distribution and are generally best characterized by the Poisson or negative binomial distributions. The modeling of both the simulated and empirical MRFSS catch rates indicates that one may draw erroneous conclusions about stock trends by assuming the wrong error distribution in procedures used to developed standardized indices of stock abundance. The results demonstrate the importance of considering not only the overall model fit and significance of classification effects, but also the possible effects of model misspecification, when determining the most appropriate model construction.

Age and growth of the bastard grunt (Pomadasys incisus: Haemulidae) inhabiting the Canarian archipelago, Northwest Africa

The bastard grunt (Pomadasys incisus) is one of the most abundant coastal demersal fishes inhabiting the Canary Islands. Age and growth were studied from samples collected between October 2000 and September 2001. Growth analysis revealed that this species is a fast growing and moderately short-lived species (ages up to seven years recorded). Length-at-age was described by the von Bertalanffy growth model (L∞=309.58 mm; k=0.220/year; t0=–1.865 year), the Schnute growth model (y1=126.66 mm; y2=293.50 mm; a=–0.426; b= 5.963), and the seasonalized von Bertalanffy growth model (L∞=309.93 mm; k=0.218/ year; t0= –1.896 year; C=0.555; ts=0.652). Individuals grow quickly in their first year, attaining approximately 60% of their maximum length; after the first year, their growth rate drops rapidly as energy is probably diverted to reproduction. The parameters of the von Bertalanffy weight growth curve were W∞=788.22 mm; k=0.1567/year; t0= –1.984 year. Fish total length and otolith radius were closely correlated, r2=0.912. A power relationship was estimated between the total length and the otolith radius (a=49.93; ν=0.851). A year’s growth was represented by an opaque and hyaline (translucent) zone—an annulus. Backcalculated lengths were similar to those predicted by the growth models. Growth parameters estimated from the backcalculated sizes at age were L∞=315.23 mm; k=0.217/year; and t0= –1.73 year.