Age and growth of the blackchin guitarfish Glaucostegus cemiculus (Geoffroy Saint-Hilaire, 1817) from Iskenderun Bay (Northeastern Mediterranean)

Between September 2010 and June 2012, a total 291 (166 females and 125 males) blackchin guitarfish Glaucostegus cemiculus were captured by a commercial bottom trawler (F/V Coşkun Reis) in Iskenderun Bay, Turkey (northeastern Mediterranean Sea). The total length (L) and total weight (W) of the female and male guitarfish ranged between 32.0–165.0 cm and 88 g–16.68 kg, and 34.3–128.3 cm and 112 g–6.00 kg, respectively. Vertebral age estimates ranged from 0 to 8 years for females and 0 to 5 years for males. The growth models of von Bertanlanffy and Gompertz were fitted to the length at age data using the nonlinear regression method. Model selection was based on the values of the residual standard error and the Akaike's information criterion corrected for small sample size (AIC_C) associated with each fit. The von Bertalanffy growth model provided the best fitting growth curves for each sex with parameters reaching L_∞ = 187.17 cm, K = 0.195 year^-1, t₀ = −1.38 year for females, and L_∞ = 144.85 cm, K = 0.321 year^-1, t₀ = −1.13 year^-1 for males. The W–L relationship parameters did not differ significantly between sexes, the estimated values of a and b were 0.0018 and 3.11, respectively. By using these values of a and b, and also respective estimates of L_∞, the values of W_∞ were obtained as 20.53 kg for females and 9.25 kg for males. The overall percentage ratios of females and males in the samples were 57% and 43% respectively.     

Age and Growth of the Whiskery Shark, Furgaleus macki, from Southwestern Australia

Age and growth of the whiskery shark, Furgaleus macki, from southwestern Australia were examined using vertebral ageing and tag-recapture data. The readability of bands on the vertebral centra varied markedly between individuals. Four readers were used to make band counts, with the most experienced reader having the lowest index of average percent error and the highest level of agreement with final counts. Marginal increment analysis indicated that opaque bands form in January. With parturition occurring from August to October, size data suggests that the first band is probably formed 15–17 months after birth. The age at maturity was estimated to be 4.5 years for males, and 6.5 years for females. The oldest male was 10.5 years, and oldest female was 11.5 years. Von Bertalanffy growth parameters for males were L =121.5cm fork length, K=0.423 year^–1, t ₀=–0.472 years, were L =120.7cm fork length, K=0.369 year^–1, t ₀=–0.544 years for females, and were L =118.1cm fork length, K=0.420 year^–1, t ₀=–0.491 years for combined sexes. Data from a tag recapture study were analysed using a maximum likelihood method to verify the estimates of growth parameters from vertebral ageing. Von Bertalanffy growth parameters from the tag recapture study were L =128.2cm fork length, K=0.288 year^–1, t ₀=–0.654 years. The two methods of estimating growth parameters produced similar results, with rapid growth until approximately 5 years of age, after which there was little increase in length.     

Reproduction,growth and feeding habits of stout beardfish Polymixia nobilis (Polymixiidae) off the Canary Islands (NE Atlantic)

The present study provides fisheries biology knowledge which will allow the implementation of regulatory measures contributing to the sustainability of the fisheries and the conservation of the stout beardfish Polymixia nobilis Lowe, 1838 off the Canary Islands, north eastern Atlantic Ocean. Males ranged between 16.5 and 38.4 cm fork length (FL) and females from 14.2 to 46.5 cm FL. Sex ratio by size classes provided significant differences in classes higher than 36 cm, being clearly unbalanced in favour of females. Individuals in maturing and mature stages were present during all months sampled, although a spawning peak is evident between April and June. Size at first maturity was estimated as 26 cm FL for females and 30 cm FL for males. Age was determined from annuli in whole otoliths. Age range was found to be 0–14 years for fish measuring 14.2 to 46.5 cm FL. It is a slow‐growing and long‐lived species. Significant differences in the growth parameters between sexes were detected. The von Bertalanffy growth parameters estimated for females (n = 213) were L_∞ = 45.92 cm LF, k = 0.16 years^?1 and t₀ = ?2.84 years; and for males (n = 186) L_∞ = 36.44 cm LF, k = 0.26 years^?1 and t₀ = ?2.16 years. Stomach analysis indicated some variations in the feeding habits with growth: individuals of small and medium sizes preyed on crustaceans and fishes, while large specimens preyed mainly on fishes.     

Age,growth, reproduction and feeding of John Dory,Zeus faber (Pisces: Zeidae), in the Saros Bay (North Aegean Sea)

The age, growth, reproduction, sexual maturity and stomach content of John Dory (Zeus faber), caught in the Bay of Saros (North Aegean Sea) between September 2006 and September 2008, were investigated. The female‐male ratio was 1.6 : 1. The total length of females ranged from 13.8 cm to 52.8 cm, and of males from 12.2 cm to 42.9 cm. The length‐weight relationship was W = 0.0174*L^2.936. Age data derived from vertebral centra readings were used to estimate the growth parameters of the von Bertalanffy equation. Growth parameters for females were L_∞ = 58.50 cm K = 0.11 year^?1, t₀ = ?0.99 year and for males L_∞ = 45.01 cm, K = 0.13 year^?1, t₀ = ?1.17 year. Maximum age was 18 years for females and 17 years for males. Length at first maturity for both females and males was 25.4 cm. Monthly values of the gonadosomatic index indicated that spawning occurred mainly in two peaks: one between January and June, and the other from August to September. Stomach content analysis showed the most‐preferred prey to be Pisces (IRI% 97.3).     

Reproduction,growth and mortality of the exploited sillaginid,Sillago ciliata Cuvier, 1829

The goal of this study was to examine the age and size composition, growth, reproductive biology and mortality of Sillago ciliata Cuvier, 1829 in one of the largest estuarine commercial fisheries in south‐eastern Australia. The study also aimed to present a qualitative comparison of latitudinal variations in some of these characteristics along the eastern Australian coastline. The sampled population contained fish aged up to 10 years with a maximum size of 39.2 cm fork length (L_F), and was dominated by 1–5 year olds. Sexual divergence in both the age and size structure of the population was recorded. Female S. ciliata grew slightly faster and attained a greater maximum size (L_∞ = 33.79 cm L_F, k = 0.50 year^?1 and t₀ = ?0.57 years) than males (L_∞ = 29.73 cm L_F, k = 0.49 year^?1 and t₀ = ?0.67 years). Females also matured at a significantly larger size (19.13 cm) and older age (1.63 years) than males (size: 17.07 cm, age: 1.10 years). Reproductive activity was highest between September and March. There were no differences between males and females in terms of mortality rate; the estimated total population, natural and fishing mortality rates were Z = 0.64, M = 0.42 and F = 0.22, respectively. Although these mortality rates suggest that S. ciliata in the Clarence River are relatively resilient to current rates of exploitation, regular monitoring of their commercial and recreational catch as well as their population structure is recommended in order to maintain sustainable fisheries. Potential latitudinal shifts in the spawning period, age structure and growth of S. ciliata along eastern Australia were also revealed.     

Age and growth of Gerres oyena (Forsskål, 1775) on Okinawa Island, Japan

Age and growth were studied for Gerres oyena (Forsskål, 1775) on Okinawa Island, southern Japan from November 2002 to November 2005. A total of 408 samples was collected ranging from 5.85 to 19.65 cm standard length (L_SL). Male fish age was estimated at up to 6⁺ years, whereas females reached 8⁺ years as estimated by sectioned otoliths. The length–weight relationships and the von Bertalanffy growth curve were described for all individuals as: and L_t = 20.54{1 − e^{−0.1807(t+2.8462)}}, respectively. Opaque rings were formed from April to August during the spring–summer seasons.     

Age and growth validation of the small spotted grunt Pomadasys commersonnii (Lacepède, 1801) from the northwestern coast of the Arabian Sea of Oman

Age structure and growth parameters were determined for a population of small spotted grunt, Pomadasys commersonnii in the Arabian Sea. Small spotted grunt samples were collected monthly between September 2007 and August 2008 by beam trawl [40 mm cod‐end mesh size] surveys conducted along the Arabian Sea coast of Oman from Ras Al‐Had in the north to the Oman‐Yemen border in the south west of Salalah between latitudes 16^o 33′N and 22^o 21′N and longitude 53^o 09′ and 59^o 55′ E and from depths of 20 to 250 m respectively. Marginal increment analysis of the sagittal otolith confirmed the deposition of annual increments on transverse sections, validating this technique for age and growth studies. Fish size (Fork Length, FL) ranged between 33–78 cm and age of fish ranged between 2 and 14 years with 5–9 years old fish comprising the majority of the catch. Both males and female exhibited asymptotic patterns of growth and the combined growth curve for both sexes provided von Bertalanffy growth (VBG) parameters of L_∞ = 77.2 cm FL (≡83.8 cm Total Length), k = 0.232/y and t₀ = ?0.058 y. VBG parameters, derived from Length‐Frequency Distribution Analysis (LFA) using ELEFAN1, PROJMAT and SLCA, obtained similar values (average values for the 3 methods: L_∞ = 78.8 cm FL, k = 0.35/y and t₀ = ?0.64 y) to those obtained from otolith analysis. A strong linear relationship was observed between otolith weight and age, and although age residuals ±4 years was observed between sagittal otolith age and estimated age (depending on sex and otolith weight), the results indicated that otolith weight could be used as a rapid proxy to estimate age and derive VBG parameters. In conclusion, both otolith ageing and LFA methods provided similar L_∞ and k values and the average values derived from all 4 methods lies within the auximetric plot for the species providing confidence in the VBG parameters obtained for small spotted grunt in the Arabian Sea.     

Growth of pike Esox lucius L. in Lake Rubikiai (Lithuania)

The aim of the study was to determine the growth rate and growth parameters of pike Esox lucius L. in the mesotrophic Lake Rubikiai and to evaluate species growth in light of environmental conditions in Lithuanian lakes of different limnologic types. A total of 204 pike specimens were caught in two subsequent years, April 2005 and April 2006, in Lake Rubikiai. Six age classes (2–7 years of age) for males and 10 age classes (3–12 years of age) for females were identified by scale readings; 2‐ to 4‐year‐old males and 4‐ to 7‐year‐old females prevailed. Total length of pike ranged from 26.5 to 107 cm and weight from 106 to 7600 g. There were no statistically significant differences (Student’s t‐test, P > 0.1) between mean lengths and weights of males and females of the same age. Von Bertalanffy growth parameters for the entire population were: L_∞ = 131.7 cm, k = 0.1530 year^?1, t_o = ?0.0398 year. Calculated overall growth performance index ?′ was 3.42. The length‐weight relationship was described by the equation . The growth rate of pike in Lake Rubikiai was higher than in most other investigated Lithuanian lakes. Pike from Lake Rubikiai are characterized by average growth, compared to other populations from various geographical locations. No significant differences were established between the growth of pike in mesotrophic and eutrophic lakes. Pike growth rate did not correlate with lake surface area, maximum or mean depth, and negatively correlated with bogginess in mesotrophic lakes and limpidity and bogginess in eutrophic lakes (P < 0.05).     

Growth characteristics of Uranoscopus scaber Linnaeus 1758, inhabiting the Iskenderun Bay (Northeastern Mediterranean)

This study aimed to determine the age and some growth characteristics of Atlantic stargazer (Uranoscopus scaber) from Iskenderun Bay (Northeastern Mediterranean). For this purpose, a total of 150 Atlantic stargazer ranging in size from 9.1 to 28.0 cm in total length (weight: 11.7–345.7 g) were collected as by-catch from a commercial trawl fishing boat at a depth of 80–100 m between May 2015 and January 2016. The bottom trawl gear used was equipped with a 44 mm stretched mesh size net at the cod-end. The percentage of females and males were 46.7% and 53.3% respectively. The total length–weight relationships equation with coefficient of determination (R²) were found as W = 0.011*TL^3.131, R² = .9728, for all individuals, W = 0.015*TL^3.021, R² = .9512 for females and W = 0.0102*TL^3.136, R² = .9553 for males. By using the von Bertalanffy equation, the growth parameters of Atlantic stargazer were estimated to be L_∞ = 42.35 cm, k = 0.098, t₀ = −1.8474 for all individuals; L_∞ = 36.92 cm, k = 0.138, t₀ = −1.2693 for females and L_∞ = 38.77 cm, k = 0.1, t₀ = −2.334 for males. In this study, age reading was done by two independent readers and index of average percentage error (IAPE) was calculated as 6.1%. The highest condition factor value calculated as 1.81 in the age group 6 and the lowest condition factor value was calculated as 1.48 in the age group 1.     

Age,growth and mortality of common two‐banded seabream,Diplodus vulgaris (Geoffroy Saint‐Hilaire, 1817), in the eastern Adriatic Sea (Croatian coast)

Age, growth and mortality were analysed for the common two‐banded seabream, Diplodus vulgaris, collected in the eastern Adriatic (Croatian coast) from commercial fishery catches by ‘tramata’ fishing (2005–2006) to obtain growth estimation. The oldest female was estimated to be age 11, the oldest male age 10 years. The von Bertalanffy growth parameters estimated by reading scales were: L_∞ = 48.60 cm (SE = 1.101), K = 0.112 (SE = 0.005) and t₀ = ?2.366 (SE = 0.060) for all specimens; L_∞ = 51.96 cm (SE = 2.153), K = 0.095 (SE = 0.007) and t₀ = ?2.837 (SE = 0.120) for females and L_∞ = 56.25 cm (SE = 2.662), K = 0.084 (SE = 0.067) and t₀ = ?2.920 (SE = 0.117) for males. The overall sex ratio was 1.22 : 1 in favour of males. Total mortality, corresponding to the slope of the descending limb of the catch curve, was Z = 0.81 per year for females and Z = 0.85 per year for males. Exploitation ratios were E = 0.68 for females and E = 0.73 for males.     

Population parameters of Mullus barbatus in Izmir Bay (Aegean Sea), using length frequency analysis

The growth and mortality of Mullus barbatus from Izmir Bay (Aegean Sea) was investigated based on length frequency data using the fisat software. Total length of the sampled 110 891 fish ranged from 5.0 to 23.0 cm. The seasonalized von Bertalanffy growth resulted in parameter values of L_∞ = 24.26 cm, k = 0.565, t₀ = ?0.305, C = 0.27 and WP = 0.45. Potential lifespan was estimated to be 5.3 years. Total mortality (Z) based on the length converted catch curve was 3.70 (seasonal data) and 3.85 (non‐seasonal data). Natural mortality (M) was 1.07 and the exploitation ratio (0.71–0.72) revealed an overfished stock.     

Length–weight relationship,parameters of growth and mortality for the hardhead halfbeak Chriodorus atherinoides Goode & Bean, 1882 (Beloniformes: Hemiramphidae) from Yucatan Peninsula,Mexico

This study reports length–weight relationships and growth parameters for Chriodorus atherinoides from La Carbonera, a karstic tropical coastal lagoon on the northwestern coast of the Yucatan Peninsula, Mexico. A total of 667 specimens were collected between April 2009 and March 2010. The length‐weight relationship obtained was W = 0.0034Lt^3.22. This study presents the first estimation of the model parameters of the von Bertalanffy growth equation (L_∞ = 27.30 cm, k = 0.76 year^?1 and t₀ = ?0.178 years), the instantaneous total mortality (2.51 year^?1) and the maximum length record (26.0 cm total length), which is greater than previously recorded.     

Growth and length–weight relationships of Sander lucioperca (Linnaeus, 1758) in the Alcántara Reservoir,south‐western Spain: comparison with other water bodies in Eurasia

Growth parameters in pikeperch may be affected by several factors. The purpose of this study was to investigate the growth and length–weight of pikeperch in a reservoir in south‐western Spain and compare the results with previous published data. Age and growth are described for Sander lucioperca from the Alcántara Reservoir (south‐western Spain) from March to October 2009. A total of 285 fish were examined; the ratio of males to females was estimated as 0.78:1 and age ranges as 1–5 years. Total lengths ranged from 16.1 to 52.5 cm. The length–weight relationships were described as W = 0.00 462 TL^3.09 (r = 0.9865) for males and W = 0.00279 TL^3.16 (r = 0.9921) for females. Growth was expressed in length and the von Bertalanffy growth parameters were estimated as L∞ = 92.14 cm, k = 0.09, t₀ = ?1.05 for males and L∞ = 107.72 cm, k = 0.08, t₀ = ?1.16 for females. Growth performance indexes were also estimated as Φ’ = 2.88 for males and Φ’ = 2.96 for females. Differences in growth and length–weight relationships between sexes were not found.     

Age,growth and mortality of scaldfish (Arnoglossus laterna Walbaum, 1792) from the Adriatic Sea

The population dynamics parameters of scaldfish, Arnoglossus laterna, from the Adriatic Sea were studied. Specimens ranging from 4.5 to 17.1 cm in total length were collected from commercial bottom‐trawl catches (2000–2006). Spawning from April through the end of August with a peak in May, A. laterna is one of the non‐target species usually present in mixed bottom trawl catches. The overall male to female ratio was 1.00 : 0.61. The oldest individual was 6 years of age. Length–weight relationships showed positive allometric growth (b = 3.36 for males and b = 3.39 for females). Parameters of the von Bertalanffy growth equation were: L_∞ = 17.3 cm; K = 0.258 per year; t₀ = ?1.0026 year; R² = 0.925. The relatively low value of total mortality (Z = 0.90) and relatively high value of constant rate of natural mortality (M = 0.64) was estimated. Presumed calculated fishing mortality (F = 0.26) and the consequently estimated exploitation ratio (E = 0.29) indicated no danger of an over‐exploitation of this species in the study area at the current level of fishing pressure. The low T max and correspondingly high M suggest that this small benthic predator plays an important role in the Adriatic ecosystem as a prey species for large benthic predators. However, the estimates include uncertainties and require further confirmation, especially with regard to natural mortality.     

Age, growth and mortality of brown comber, Serranus hepatus (Linnaeus, 1758) (Pisces: Serranidae), in the eastern Adriatic (Croatian coast)

Age, growth and mortality were analysed for brown comber, Serranus hepatus, collected in the eastern Adriatic from May 2002 to June 2003. Fish aged 2–7 years were present in the samples. Growth in length including both sexes was expressed using the von Bertalanffy equation, L_∞ = 14.82, K = 0.217 and t₀ = −1.672. Age class 2, ranging in length from 5.8 to 9.0 cm TL, dominated the sample (29.3%). Total and natural mortality were Z = 1.73 year⁻¹ and M = 0.63 year⁻¹, respectively. The exploitation rate, E = 0.64, revealed a high fishing pressure on the stock in the studied area.     

Age,growth and mortality of Lutjanus alexandrei in estuarine and coastal waters of the tropical south‐western Atlantic

Otolith‐based methods were used to determine life history traits of the endemic Brazilian snapper (Lutjanus alexandrei) in estuarine and coastal environments in the south‐western Atlantic. Fishes were caught as juveniles inside mangrove‐bordered estuaries by traditional corral fisheries whereas adults were captured at sea using motorboats with trap and gill nets. Fish were sampled during landings and 331 otolith pairs were extracted from L. alexandrei. Inshore mangroves comprised individuals of 0–4 years (mean: 2 years), while individuals in deeper reef environments were older (range: 3–22; mean: 8 years), indicating an ontogenetic shift at approximately age 3 or 4. Edge analysis was used to validate the annual deposition in the otoliths, suggesting that opaque growth rings were formed between April and September. Age‐at‐length data were used to predict L. alexandrei growth rates using the von Bertalanffy growth model from where the parameters were calculated: L_∞ = 31 cm, k = 0.24, t₀ = ?1.26, r² = 0.97. Mortality rates were estimated for coastal habitats, with Z = 0.22 and S = 0.78 year^?1, based on ages 7–17. Additionally, evidence of ontogenetic migration is provided by age and size structure.     

Age,growth, and age at sexual maturity of the commercially landed skate species,Dipturus chinensis (Basilewsky, 1855), in the northern East China Sea

Age, growth, and age at sexual maturity of the polkadot skate Dipturus chinensis, in the northern East China Sea were determined for a total of 614 specimens collected from April 2009 to December 2014. Vertebral centrum analysis was used to calculate the age of the skates. Annual band deposition was determined by marginal increment analysis. The von Bertalanffy growth model was fitted to the observed length‐at‐age data for each sex (males, L_∞ = 76.8, k = 0.109, t₀ = ?1.28; females, L_∞ = 83.1, k = 0.103, t₀ = ?1.20). Growth patterns of females and males were similar until the age of 6; thereafter, females grew larger than males. Maximum age recorded was 13 years for males and 15 years for females. Age at 50% sexual maturity was 8.22 years for males and 9.39 years for females. These results indicate that D. chinensis is slow growing, relatively long‐lived, and late maturing, and therefore vulnerable to exploitation.     

Geographic variation in life-history traits of the endemic kite skate Dipturus chilensis (Batoidea: Rajidae), along its distribution in the fjords and channels of southern Chile

Life-history traits of kite skates Dipturus chilensis were examined from two regions (c. 2286 km apart) in the sheltered fjords and channels of southern Chile. A total of 482 and 403 specimens were collected from the southern fjords (c. 42–46° S) and the fjords of Chilean Patagonia (c. 51–54° S), from September 2003 to 2004, respectively. Vertebra marginal increment analysis indicated an annual deposition of growth rings which was completed during the winter months. For each region, von Bertalanffy growth parameters showed that females attained a larger asymptotic size, L_∞, had a lower growth coefficient, K, and lived longer than males. Growth analysis indicated that D. chilensis from the Patagonian fjords had a longer life span (females: 22 v. 21 years; males 19 v. 17 years), attained a larger L_∞ (females: 150 v. 136 cm; males: 122 v. 118 cm total length, L_T) and had a lower K value (females: 0·087 v. 0·104; males: 0·110 v. 0·116) than their counterparts in the southern fjords. Comparisons with previous studies indicated that D. chilensis from both southern and Patagonian sheltered fjords had larger L_∞, and grew more slowly than their counterparts from central-southern Chile (L_∞= 119–123 cm, K= 0·123–0·127), suggesting latitudinal variations in growth. Females attained sexual maturity later than males in both regions. For both sexes, lengths at 50% maturity (L_50%) between regions were similar (females: c. 103 cm; males: c. 87 cm L_T); however, D. chilensis from Patagonia appeared to mature 1 year earlier (females: 13 v. 14 years; males: 10 v. 11 years). Specimens from Patagonia had a lower ovarian fecundity than those from the southern fjords. An increase in the proportion of mature females and males during summer, suggests that the reproductive peak occurs in this season, and no regional differences were found. The size of the egg cases increased with maternal L_T and these were longer in Patagonia. The information provided here represents the first evidence of regional variations in life-history traits for elasmobranchs in the south-eastern Pacific.     

Age and growth of the hollowsnout grenadier, Caelorinchus caelorhincus (Risso, 1810), in the Aegean Sea

Age and growth of the hollowsnout grenadier was studied based on samples collected seasonally in the Agean Sea between March 2003 and January 2004. Total length ranged from 9.0 to 21.6 cm. Length–weight relationship was described as a = 0.0032, and b = 3.008. Age determinations based on otolith readings were between 1 and 10 years. The von Bertalanffy growth curve fitted to observed lengths‐at‐age provided parameters of L_∞ = 24.9 cm, k = 0.115 and t_o = ?3.494.     

Population parameters of stargazer (Uranoscopus scaber Linnaeus, 1758) in the southeastern Black Sea region during the 2011–2012 fishing season

The population parameters of stargazer (Uranoscopus scaber Linnaeus, 1758) were studied regarding age composition, sex ratio, growth, survival and mortality rates, and the exploitation rate in the southeastern Black Sea near the coast of Turkey during the 2011–2012 fishing season. According to ageing analysis of the samples, age group 1 is the most abundant (47.76%), followed by age 2 (41.04%), age 3 (9.33%), age 0 (1.12%) and age 4 (0.75%). Mean totals as well as total lengths and weights for males and females were 14.5 ± 0.22 cm and 12.9 ± 0.19 g, 16.6 ± 0.24 cm and 62.1 ± 2.76 g, and 37.2 ± 1.92 cm and 87.1 ± 3.88 g, respectively. The mean condition factor was K = 0.0167, while the sex ratio was 53.98% female, 38.75% male and 7.27% immature. Length‐weight, age‐length and age‐weight equations were W = 0.014 × L^3.059, L_(t)=44.5*[1?e^{?0.148*(t +1.242)}] and W_(t)=1544.4*[1?e^{?0.148*(t + 1.242)}]^3.059, respectively. The computed survival rate (S), instantaneous total mortality rate (Z), annual mortality rate (A), natural mortality rate (M) and fishing mortality rate (F) were S = 28.94%, Z = 1.24 year^?1, A = 71.06%, M = 0.26 year^?1 and F = 0.98 year^?1, respectively. The exploitation rate was 0.79, which is above the optimum exploitation levels.