Economics of harvesting age-structured fish populations

A generic age-structured model is developed to derive analytical results on optimal harvesting. Given two age classes, knife-edge selectivity, and no stock-dependent harvesting cost, the steady state is a unique saddle point. Adding harvesting cost does not alter the uniqueness, given that the utility is linear. Under specific conditions such as nonselective gear, optimal harvesting is proved to be a stationary cycle that represents pulse fishing. Optimal steady states are different if age-structured information is ignored and optimization is based on traditional biomass variables. This implies that the existence of optimal sustainable harvesting depends on age-structured information. Given a specific set of conditions such as low interest rate and knife-edge selectivity, optimal harvesting converges toward a unique saddle point independently of the number of age classes.     

Economics of harvesting age-structured fish populations

A generic age-structured model is developed to derive analytical results on optimal harvesting. Given two age classes, knife-edge selectivity, and no stock-dependent harvesting cost, the steady state is a unique saddle point. Adding harvesting cost does not alter the uniqueness, given that the utility is linear. Under specific conditions such as nonselective gear, optimal harvesting is proved to be a stationary cycle that represents pulse fishing. Optimal steady states are different if age-structured information is ignored and optimization is based on traditional biomass variables. This implies that the existence of optimal sustainable harvesting depends on age-structured information. Given a specific set of conditions such as low interest rate and knife-edge selectivity, optimal harvesting converges toward a unique saddle point independently of the number of age classes.     

Optimal harvesting of ecologically interdependent fish species

The optimal exploitation of a two-species predator-prey system is considered, using Lotka-Volterra-type equations. Due to the density-dependence of ecological efficiency, both species should be harvested simultaneously over a range of relative prices. Beyond the limits of this price range, either the prey species should be utilized indirectly by harvesting the predator, or the predator should be eliminated to maximize the prey yield. Neglecting harvesting costs, the simultaneous harvest of prey and predators requires that a unit of prey biomass increase in value by being “processed” by predators. Certain results from single-species fishery models are shown not to apply to multispecies models. These are as follows: (i) Optimal regulation of a free access fishery may call for subsidizing instead of taxing the harvest of predator species. (ii) Increasing the discount rate may, at “moderate” levels, imply that the optimal standing stock of biomass increases instead of decreasing. (iii) A rising price or a falling cost per unit fishing effort of a species may raise and not lower the optimal standing stock of that species.     

The energetic cost of protogynous versus protandrous sex change in the bi-directional sex-changing fish Gobiodon histrio

To investigate the relative cost of protogynous versus protandrous sex change we induced sex change in each direction in Gobiodon histrio (Gobiidae) and then compared growth, body condition and biochemical condition between sex-changed and non-sex-changed fish in treatment and control groups. Sex change in each direction was induced by establishing pairs of adult males and pairs of adult females on isolated coral colonies. Heterosexual pairs were used as controls. For both males and females, growth and body condition did not vary between fish that changed sex and those that did not. The relatively low cost of sex change, compared to the likely costs of searching for a mate of the correct sex, appears to explain the evolution of bi-directional sex change in coral-dwelling gobies. Lipid concentrations in the liver of males and females that changed sex were reduced by similar amounts (34% and 41%, respectively) compared to male and female controls that did not change sex. Therefore, changes in biochemical condition were approximately equal for fish that sex change in each direction and cannot explain the unequal frequency of sex change in each direction in natural populations of G. histrio.     

Selective harvesting and habitat loss produce long-term life history changes in a mouflon population.

We examined the long-term effects (28 years) of habitat loss and phenotype-based selective harvest on body mass, horn size, and horn shape of mouflon (Ovis gmelini musimon) in southern France. This population has experienced habitat deterioration (loss of 50.8% of open area) since its introduction in 1956 and unrestricted selective hunting of the largest horned males since 1973. Both processes are predicted to lead to a decrease in phenotype quality by decreasing habitat quality and by reducing the reproductive contribution of individuals carrying traits that are targeted by hunters. Body mass and body size of both sexes and horn measurements of males markedly decreased (by 3.4-38.3%) in all age classes from the 1970s. Lamb body mass varied in relation to the spatiotemporal variation of habitat closure within the hunting-free reserve, suggesting that habitat closure explains part of these changes. However, the fact that there was no significant spatial variation in body mass in the early part of the study, when a decline in phenotypic quality already had occurred, provided support for the influence of selective harvesting. We also found that the allometric relationship between horn breadth and horn length changed over the study period. For a given horn length, horn breadth was lower during the second part of the study. This result, as well as changes in horn curve diameter, supports the interpretation that selective harvesting of males based on their horn configuration had evolutionary consequences for horn shape, since this phenotypic trait is less likely to be affected by changes in habitat characteristics. Moreover, males required more time (approximately four years) to develop a desirable trophy, suggesting that trophy hunting favors the reproductive contribution of animals with slow-growing horns. Managers should exploit hunters' desire for trophy males to finance management strategies which ensure a balance between the population and its environment. However, for long-term sustainable exploitation, harvest strategy should also ensure that selectively targeted males are allowed to contribute genetically to the next generations.     

Contrasting demographic histories in a pair of allopatric,sibling species of fish (Arripidae) from environments with contrasting glacial histories

Although Pleistocene glacial cycles are regularly used to explain many aspects of the demographic history of coastal marine species, the present study is one of few to test a priori predictions about these effects. Nucleotide sequence variation in the cytochrome b gene of the mtDNA and allele frequency variation at intron loci in the nDNA were compared between two species of Arripis (Australian Salmon) that are closely related and biologically similar other than for their allopatric distributions. The results suggest that A. truttaceus, which resides to the west of the Bass Strait, exhibits lower levels of genetic diversity and experienced a severe population bottleneck during the LGM followed by an expansion commencing some 17,000 years ago. In contrast, the population of A. trutta, which resides to the east of the Bass Strait, appears to have been largely unaffected by the LGM and has been expanding over the past 100,000 years or more. These results are consistent with a priori predictions, based on paleo-oceanographic data, that the demographic history of A. truttaceus has been more strongly affected by glacial periods by virtue of its distribution to the west of Bass Strait. Data on two other congeners are also presented to provide context for the results for A. trutta and A. truttaceus.     

Vertical structure of very nearshore larval fish assemblages in a temperate rocky coast

Small-scale vertical patterns of larval distribution were studied at a very nearshore larval fish assemblage, during the spring–summer period of several years, at two depth strata (surface and bottom) using sub-surface and bottom trawls. A total of 4,589 larvae (2,016 from surface samples and 2,573 from bottom samples) belonging to 62 taxa included in 22 families were collected. Most larvae belonged to coastal species. Although inter-annual variations in larval density and diversity could be found, total larval abundance was always higher near the bottom whereas diversity was higher at the surface. A marked distinction between the structure of surface and bottom assemblages was found. Sixteen taxa explained 95% of the similarity among surface samples. Larvae which contributed most to this similarity included species like clupeiformes, sparids and serranids, and also blenniids, tripterygiids and some labrids. In the bottom samples, fewer species were present, with only six taxa, almost exclusively from species which lay demersal eggs, contributing to 95% of the similarity between samples. Larvae present at the surface were significantly smaller than at the bottom. For some of the most abundant species found at the bottom, only small larvae occurred at the surface while the whole range of sizes was present at the bottom, indicating that larvae may be completing the entire pelagic phase near the adults’ habitat. These results indicate that larval retention near the reefs probably occurs for these species, although for others dispersal seems to be the prevailing mechanism.     

Pre-reproductive survival in a tropical bird and its implications for avian life histories

The factors that affect survival until reproduction are essential to understanding the organization of life histories within and among species. Theory predicts, for example, that survival until reproduction influences the optimum level of reproductive investment by parents, which might partly explain prolonged parental care in species with high first-year survival. Tests and refinements of life-history theory have been hampered, however, by a lack of field-based estimates of pre-reproductive survival, especially for tropical species, which have been the subject of many comparative analyses. Tropical species are predicted to have higher first-year survival and delayed reproduction compared to Northern Hemisphere species. We estimated survival until reproduction, age at first reproduction, and sources of variation in juvenile survival in a Neotropical passerine, the Western Slaty-Antshrike (Thamnophilus atrinucha), in central Panama. We observed that fledged antshrikes had 76% survival through the dependent period and 48% survival to the age of 1 year; survival rate was lowest during the first week after leaving the nest. Timing of fledging within the breeding season, fledgling mass, and age at dispersal influenced survival, while sex of offspring and year did not. Individuals did not breed until two years of age, and post-fledging pre-reproductive survival was 41% of annual adult survival. High survival until reproduction in antshrikes balanced their low annual productivity, resulting in a stable population. Survival during the post-fledging period of dependence and the first year of independence in the Western Slaty-Antshrike exceeded estimates for Northern Hemisphere species. This difference appears to be associated with the extended post-fledging parental care, delayed dispersal, low costs of dispersal, and the less seasonal environment of antshrikes.     

Home range, activity and distribution patterns of a temperate rocky-reef fish, Cheilodactylus fuscus

Red morwong (Cheilodactylus fuscus; Cheilodactylidae: Perciformes), are large temperate rocky reef fish that are patchily distributed in local aggregations of 3 to >100 fish. The home ranges and aggregating behaviour of red morwong were investigated using external tags (n = 114, over 35 mo) and ultrasonic transmitters (n = 9 over 20 d) at coastal and estuarine sites in south-east Australia. Adult red morwong had a mean home range during the day of 1865 m² (SE = 268). Night movement, determined by ultrasonic telemetry, indicated a significantly greater mean home range of 3639 m² (SE = 416). Home range did not differ significantly with size or sex. Fish movements were greatest and aggregation sizes varied most during the non-reproductive mid-summer period. Tagged fish (N = 20) displaced 200 to 900 m returned in 1 to 3 d to the point of capture, often traversing open sand habitat and other aggregations. During the day, fish were commonly found on bouldered habitat. During the night, these site-specific aggregations fragmented as fish dispersed over a variety of substrata, with crepuscular peaks in activity. Therefore, estimates of habitat-specificity and feeding patterns collected only by day may give misleading results. Received: 28 June 1996 / Accepted: 8 July 1998     

The sensitivity of carbon sequestration to harvesting and climate conditions in a temperate cypress forest: Observations and modeling

The role of disturbance and climate factors in determining the forest carbon balance was investigated at a Japanese cypress forest in central Japan with eddy flux measurements, tree-ring analyses, and a terrestrial biosphere model. The forest was established as a plantation after intermittent harvesting and replanting between 1959 and 1977, and acted as a strong carbon sink of approximately 500 g C m⁻² year⁻¹ for the measurement years between 2001 and 2007. A terrestrial biosphere model, BIOME-BGC, was validated using the eddy flux data at daily to interannual timescales, and the tree-ring width data at interannual to decadal timescales. According to the model simulation, during the observation period 270 ± 55 g C m⁻² year⁻¹ was additionally sequestered due to the indirect effects of the harvesting and planting, whereas the increase of CO₂ concentration and the change in climate increased the sink of 110 ± 40 and 30 ± 80 g C m⁻² year⁻¹, respectively. The model simulation shows that the forest is now recovering from harvesting, and that harvesting is a more important determinant of the current carbon sink than either interannual climate anomalies or increased atmospheric CO₂ concentration. We found that harvesting with long rotation length could be effective management activity in order to increase carbon sequestration, if the harvested timber is converted into products with long lifecycles.     

The role of life histories and trophic interactions in population recovery

Factors affecting population recovery from depletion are at the focus of wildlife management. Particularly, it has been debated how life‐history characteristics might affect population recovery ability and productivity. Many exploited fish stocks have shown temporal changes towards earlier maturation and reduced adult body size, potentially owing to evolutionary responses to fishing. Whereas such life‐history changes have been widely documented, their potential role on stock's ability to recover from exploitation often remains ignored by traditional fisheries management. We used a marine ecosystem model parameterized for Southeastern Australian ecosystem to explore how changes towards “faster” life histories might affect population per capita growth rate r. We show that for most species changes towards earlier maturation during fishing have a negative effect (3–40% decrease) on r during the recovery phase. Faster juvenile growth and earlier maturation were beneficial early in life, but smaller adult body sizes reduced the lifetime reproductive output and increased adult natural mortality. However, both at intra‐ and inter‐specific level natural mortality and trophic position of the species were as important in determining r as species longevity and age of maturation, suggesting that r cannot be predicted from life‐history traits alone. Our study highlights that factors affecting population recovery ability and productivity should be explored in a multi‐species context, where both age‐specific fecundity and survival schedules are addressed simultaneously. It also suggests that contemporary life‐history changes in harvested species are unlikely to increase their resilience and recovery ability.     

Seasonal and spatial changes in the larval fish fauna within a large temperate Australian estuary

A total of 66814 fish larvae, representing 37 families and 74 species, were collected in samples taken monthly between January 1986 and April 1987 from 13 sites located at frequent intervals throughout the large Swan Estuary in south-western Australia. The Gobiidae was the most abundant family, comprising 88.2% of the total number of larvae, followed by the Clupeidae (3.4%), Engraulididae (2.9%) and Blenniidae (1.0%). The most abundant species were Pseudogobius olorum (53.3%), Arenigobius bifrenatus (31.2%) and Engraulis australis (2.9%). Abundance of fish larvae in the lower, middle and upper regions of the estuary each reached a maximum between mid-spring and early summer, 2 to 4 mo before the attainment of maximum temperatures. Larvae of species such as Nematalosa vlaminghi and Apogon rueppellii were collected only between November and February, whereas those of others such as P. olorum, E. australis and Leptatherina wallacei were present over many months. The times and locations of capture of larvae have been related to the distribution and breeding periods of the adults of these species. The mean monthly number of species was far greater in the lower than upper estuary (14.7 vs 2.7), whereas the reverse was true for mean monthly concentration (42 vs 197 larvae per 100 m³). Classification, using the abundance of each of the 74 species recorded at the different sites, showed that the composition of the larval fish fauna in the lower, middle and upper estuary differed markedly from each other. Most larvae caught in the lower estuary belonged to marine species, whereas those in the upper estuary almost exclusively represented species that spawn within the estuary. The fact that the larvae of the 59 species of marine teleosts recorded during this study were restricted mainly to the lower estuary, and yet contributed only 6.2% to the total numbers for the whole estuary, helps to account for the relatively high species diversity in this region. The lack of penetration of many of these larvae beyond the first 12.5 km of the estuary presumably reflects the weak tidal effect in the wide basins of the middle estuary and saline regions of the tributary rivers. The larvae of the 13 teleosts that typically spawn within the estuary contributed 93.8% to the total numbers of larvae. Most of these estuarine-spawned larvae belong to teleosts that deposit demersal eggs and/or exhibit parental care (egg-guarding and oral and pouch-brooding), characteristics which would maximize their chances of retention within the estuary.     

Direct versus indirect effects of wave exposure as a structuring force on temperate cryptobenthic fish assemblages

The structure of cryptic reef fish assemblages was assessed on sheltered and exposed aspects of coastal breakwaters at two locations in the northwestern Adriatic Sea. There were distinct differences between the two levels of exposure, which were consistent between locations. Habitat characteristics, measured on scales of tens of centimetres, explained 50% of the variability in assemblage structure between exposures, whereas ‘exposure’ alone (implying direct effects of wave energy on the fish) explained <5% of the variation. The most important explanatory variables were the presence of macroalgae, sandy habitat and oyster shell, the last of which increased the degree of small-scale complexity and provided nesting sites for blennies. We found little evidence to suggest that wave action had large direct effects on the fish assemblages, although this may be in part due to the relatively small degree of difference between ‘exposed’ and ‘sheltered’ samples under the calm conditions of a sea with a relatively short fetch. These results suggest that wave action acts mainly indirectly as a structuring force on cryptic reef fish communities, by altering the composition and/or the relative density of epibiota that influence the distribution of fish. Thus, relative wave energy may provide a useful means of predicting fish assemblage structure only at large spatial scales. Microhabitat, composed of a combination of physical complexity and biological elements, always explained the greater part of variability at small (<1 m) spatial scales.     

Climatic effects on the growth of a temperate reef fish from the Southern Hemisphere: a biochronological approach

Increments in the hard parts of marine organisms (otoliths, skeletons, shells) can provide long-term chronologies of growth analogous to tree rings. For the first time in the Southern Hemisphere, we use a dendrochronological (tree-ring analysis) approach to develop a multidecadal chronology of growth for a temperate reef fish, Girella tricuspidata, from the coast of northern New Zealand. Growth patterns in the otoliths of this species were strongly synchronous among individual fish over a period spanning 27 years (1980–2006). We then compared our otolith chronology to climatic records and found strong positive correlations of growth with sea surface temperature, and weak negative correlations with the multivariate El Nino Southern Oscillation (ENSO) index. Strongest correlations were found between summer sea surface temperature and otolith growth. This relationship was consistent across all years and explained 44 % of the variation (y = −2.0 + 0.1785 × temperature, r ² = 0.4367, P = 0.0002) in the G. tricuspidata growth chronology. Our study illustrates how otolith chronologies provide remarkable records of annual growth patterns over decadal time scales that will be useful for forecasting the likely effects of climate change on marine ecosystems.     

Protection from fishing alters the species composition of fish assemblages in a temperate-tropical transition zone

Closure of areas to fishing is expected to result in an increase in the abundance of targeted species; however, changes to populations of species not targeted by fishermen will depend upon their role in the ecosystem and their relationship with targeted species. The effects of protection on targeted and non-targeted reef fish species at the Houtman Abrolhos Islands, Western Australia were studied using baited remote underwater stereo–video cameras. Video images were collected from shallow (8–12 m) and deep (22–26 m) reef sites inside a Marine Protected Area (MPA) at each of three island groups and from three replicate fished locations at each of these groups that span a temperate-tropical transition area. The MPAs were established in 1994 and vary in size from 13.72 km² at the Pelsaert group in the south to 22.29 km² at the Easter group to 27.44 km² at the Wallabi group in the north. The relative abundances of 137 fish species from 42 families were recorded. Large differences in fish assemblage structure existed between MPA and fished locations, and also between shallow and deep regions. Targeted fish species Plectropomus leopardus, Lethrinus miniatus, Lethrinus nebulosus, Pagrus auratus and Glaucosoma hebraicum were more abundant inside MPAs than in areas open to fishing. Their abundance inside MPAs was between 1.13 and 8 times greater than their abundance at fished locations. For non-targeted fish species many were more abundant in areas open to fishing, e.g. Coris auricularis, Thalassoma lutescens, Thalassoma lunare, Dascyllus trimaculatus, however others were conversely more abundant inside MPAs, e.g. Gymnothorax spp, Kyphosus sydneyanus, Scarus microhinos, Chromis westaustralis, Chaetodon spp. This study demonstrates that the removal of abundant targeted species from an ecosystem by fishing can indirectly impact non-fished species and alter the trophic structure of fish assemblages. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A comparison of temperate reef fish assemblages recorded by three underwater stereo-video techniques

Three underwater stereo-video techniques were used to sample the relative densities and species richness of temperate reef fish assemblages at three reef locations and two habitats (high- and low-relief reef) within Hamelin Bay, south-western Australia. The three techniques compared were diver-operated stereo-video strip transects, baited remote stereo-video and unbaited remote stereo-video. While unbaited remote stereo-video and diver-operated stereo-video transects recorded greater species richness at high compared to low-relief reefs, baited remote stereo-video recorded similar species richness at the two habitat types. The diver-operated stereo-video system was manoeuvred through caves and under overhangs recording small, cryptic, cave-dwelling species that were not recorded by either remote video techniques (Trachinops noarlungae, Trachinops brauni, Chromis klunzingeri, Trachichthys australis). Both remote video techniques recorded greater species richness and relative density of the most common species of Labridae, Ophthalmolepsis lineolatus. Baited remote video recorded the rarer, large predatory fish species (e.g. Seriola hippos, Glaucosoma hebraicum, Heterodontus portusjacksoni). None of the techniques sampled small cryptic fish families such as Gobiidae or Blenniidae. A combination of survey techniques is recommended for comprehensive fishery-independent studies that aim to sample broad components of fish assemblages.An erratum to this article can be found at     

Settlement patterns and presettlement behavior of the naked goby,Gobiosoma bosci,a temperate oyster reef fish

Dispersion, distribution, development and feeding incidence of larvae of the naked goby,Gobiosoma bosci (Lacepéde), were examined for linkages between larval behavior while near the reef surface and later patterns of settlement and recruitment. Field sampling and experiments were conducted during the summers of 1988 and 1989 in the Flag Pond oyster reef along the western shore of the Chesapeake Bay near Camp Conoy, Maryland, USA. Results indicated that prior to settlement most demersal larvae aggregate in shoals and exhibit distinct microhabitat preferences on the reef. In a field experiment, larvae settled both during the day and at night. Dispersion at settlement was aggregated, suggesting that demersal shoaling influences settlement patterns in this species. The distribution of demersal larvae also indicated that larval swimming behavior is sufficiently strong to permit active control of position on reefs. Large demersal larvae settled rapidly when brought to the laboratory, but small larvae in demersal shoals appeared to require additional growth and morphological development prior to settlement. Development of the pelvic fins, used by juveniles and adults for perching on the substrate, may be a good indicator of competence to settle in this species. The adaptive significance of demersal shoaling by small larvae of the naked goby, and the fate of these larvae, remains perplexing because the low feeding rates found for larvae shoaling near the reef surface should slow or prevent the growth and development required prior to settlement. Observations made by other authors indicate that demersal shoaling and the use of water directly overlying reefs may be common behaviors of temperate and tropical reef fishes.     

Changes in life histories of cladocerans (Cladocera) from the rotifer-mediated allelochemicals

ABSTRACT

We hypothesised that at higher densities herbivorous rotifers through their allelochemicals affect the survival and reproduction of cladocerans. To test this, we separately cultured three rotifers (Brachionus calyciflorus, Brachionus havanaensis, and Plationus patulus) and three cladocerans (Ceriodaphnia dubia, Daphnia pulex, and Moina macrocopa). Chlorella vulgaris was used as food and reconstituted moderately hard water was used as medium. The conditioned-medium (CM) obtained from each of rotifer species was used to conduct cohort life table tests of cladocerans. SDS PAGE electrophoresis revealed the molecular weights of proteins present in the rotifer conditioned medium were 55–63?kDa. Compared to C. dubia and D. pulex, Moina was more resistant to rotifer-mediated chemicals. Gross reproductive rate of C. dubia was significantly stimulated by the rotifer-CM. However, gross and net reproductive rates of D. pulex were strongly reduced (72–85%) due to P. patulus-CM but not from the other two rotifer species. The rate of population increase of M. macrocopa was significantly reduced by the rotifer CM. Overall, about 46% (adverse effect: 24%; stimulatory impact: 22%) of the life history variables in controls significantly differed from rotifer-CM treatments, thus proving our hypothesis that rotifer-mediated allelochemicals had some effect on the cladoceran life history variables.     

Econometric modeling of fisheries with complex life histories: Avoiding biological management failures

Economics of the fishery has focused on the wastefulness of common pool resource exploitation. Pure open access fisheries dissipate economic rents and degrade biological stocks. Biologically managed fisheries also dissipate rents but are thought to hold biological stocks at desired levels. We develop and estimate an empirical bioeconomic model of the Gulf of Mexico gag fishery that questions the presumptive success of biological management. Unlike previous bioeconomic life history studies, we provide a way to circumvent calibration problems by embedding our estimation routine directly in the dynamic bioeconomic model. We nest a standard biological management model that accounts for complex life history characteristics of the gag. Biological intuition suggests that a spawning season closure will reduce fishing pressure and increase stocks, and simulations of the biological management model confirm this finding. However, simulations of the empirical bioeconomic model suggest that these intended outcomes of the spawning closure do not materialize. The behavioral response to the closure appears to be so pronounced that it offsets the restriction in allowable fishing days. Our results indicate that failure to account for fishing behavior may play an important role in fishery management failures.