Length‐ and age‐based demographics of exploited populations of stout whiting,Sillago robusta Stead, 1908

This study assessed variability in the length and age compositions, longevity, length‐at‐age and rates of growth and mortality of the east Australian stout whiting Sillago robusta Stead, 1908 population harvested by demersal trawl fisheries. Sampling was done over 2 years and was spatially stratified across three depth strata between 11 and 90 m at two locations approximately 400 km apart. There were no consistent depth‐related differences in length and age compositions, but the mean and median length and age of the population was greater at the lower latitude location. Age classes 2 and 3 years dominated samples in the north, and 1 and 2 years in the south. Observed longevity was 10 years in the north, and 6 years in the south. Mean length‐at‐age was not consistently different between sexes, years or locations, nor did the von Bertalanffy growth function differ significantly between sexes, even though females had a greater estimated L_∞ (23.45 cm FL) compared to males (22.36 cm FL). Estimated natural mortality (M) ranged between 0.42 and 0.77, using age‐ and length‐based methods. Age‐based catch‐curve analyses identified the instantaneous rate of total mortality (Z) to range between 1.48 and 2.70, with subsequent estimates of fishing mortality (F) ranging between 1.15 and 2.00, being greater than M. Exploitation rates (E) were greater than 0.7, indicating that S. robusta at the study locations was heavily fished. The data provided here can be used as a basis to evaluate future fishery‐ and climate‐related changes in the population demographics of east Australian S. robusta.     

Survival in northern microrefugia in an endemic Carpathian gammarid (Crustacea: Amphipoda)

Gammarus leopoliensis (Crustacea: Amphipoda) is considered a north‐eastern Carpathian endemic species and therefore can be regarded as an appropriate model for testing the hypothesis of Quaternary glacial survival in northern microrefugia. However, 250 km south, the south‐western Carpathians harbour populations that resemble phenotypically both G. leopoliensis and Gammarus kischineffensis, a similar species distributed east of the Carpathians. We used maximum‐likelihood and Bayesian methods to evaluate the phylogenetic relationships of these three taxa based on mitochondrial and nuclear markers, and quantitatively compared diversity patterns, phylogeography and divergence times among north‐eastern and south‐western Carpathian taxa. Results indicate that G. leopoliensis and the south‐western populations form together a strongly supported group (G. leopoliensis s.l.) which, along with G. kischineffensis, belongs to the Gammarus balcanicus clade. This group contains 12 lineages mainly of Pliocene age. G. leopoliensis consists of two widely distributed and recently expanded allopatric sister lineages that diverged from the southern ones ca. 4 Ma, indicating long‐term survival in northern microrefugia. The southern lineages are micro‐endemic and display a scattered distribution, suggesting a more ancient, relict pattern. We conclude that the contrasting diversity patterns between the disjunct distributional areas of G. leopoliensis s.l. reflect differential survival of lineages across the latitudinal gradient, offering a promising system for comparing the evolutionary ecology of lineages persisting in latitudinally disconnected microrefugia. These results fill an important gap in the knowledge of European gammarid biogeography and reveal that all Carpathian Gammarus taxa are ancient and diverse species complexes.     

Waterscape genetics of the yellow perch ( Perca flavescens): patterns across large connected ecosystems and isolated relict populations

Comparisons of a species' genetic diversity and divergence patterns across large connected populations vs. isolated relict areas provide important data for understanding potential response to global warming, habitat alterations and other perturbations. Aquatic taxa offer ideal case studies for interpreting these patterns, because their dispersal and gene flow often are constrained through narrow connectivity channels that have changed over geological time and/or from contemporary anthropogenic perturbations. Our research objective is to better understand the interplay between historic influences and modern‐day factors (fishery exploitation, stocking supplementation and habitat loss) in shaping population genetic patterns of the yellow perch Perca flavescens (Percidae: Teleostei) across its native North American range. We employ a modified landscape genetics approach, analysing sequences from the entire mitochondrial DNA control region and 15 nuclear DNA microsatellite loci of 664 spawning adults from 24 populations. Results support that perch from primary glacial refugium areas (Missourian, Mississippian and Atlantic) founded contemporary northern populations. Genetic diversity today is highest in southern (never glaciated) populations and also is appreciable in northern areas that were founded from multiple refugia. Divergence is greater among isolated populations, both north and south; the southern Gulf Coast relict populations are the most divergent, reflecting their long history of isolation. Understanding the influence of past and current waterway connections on the genetic structure of yellow perch populations may help us to assess the roles of ongoing climate change and habitat disruptions towards conserving aquatic biodiversity.     

Phylogeography of Pinus tabulaeformis Carr. (Pinaceae), a dominant species of coniferous forest in northern China

How coniferous trees in northern China changed their distribution ranges in response to Quaternary climatic oscillations remains largely unknown. Here we report a study of the phylogeography of Pinus tabulaeformis, an endemic and dominant species of coniferous forest in northern China. We examined sequence variation of maternally inherited, seed‐dispersed mitochondrial DNA (mtDNA) (nad5 intron 1 and nad4/3–4) and paternally inherited, pollen‐ and seed‐dispersed chloroplast DNA (cpDNA) (rpl16 and trnS‐trnG) within and among 30 natural populations across the entire range of the species. Six mitotypes and five chlorotypes were recovered among 291 trees surveyed. Population divergence was high for mtDNA variation (G_ST = 0.738, N_ST = 0.771) indicating low levels of seed‐based gene flow and significant phylogeographical structure (N_ST > G_ST, P < 0.05). The spatial distribution of mitotypes suggests that five distinct population groups exist in the species: one in the west comprising seven populations, a second with a north–central distribution comprising 15 populations, a third with a southern and easterly distribution comprising five populations, a fourth comprising one central and one western population, and a fifth comprising a single population located in the north‐central part of the species’ range. Each group apart from the fourth group is characterized by a distinct mitotype, with other mitotypes, if present, occurring at low frequency. It is suggested, therefore, that most members of each group apart from Group 4 are derived from ancestors that occupied different isolated refugia in a previous period of range fragmentation of the species, possibly at the time of the Last Glacial Maximum. Possible locations for these refugia are suggested. A comparison of mitotype diversity between northern and southern subgroups within the north‐central group of populations (Group 2) showed much greater uniformity in the northern part of the range both within and between populations. This could indicate a northward migration of the species from a southern refugium in this region during the postglacial period, although alternative explanations cannot be ruled out. Two chlorotypes were distributed across the geographical range of the species, resulting in lower levels of among‐population chlorotype variation. The geographical pattern of variation for all five chlorotypes provided some indication of the species surviving past glaciations in more than one refugium, although differentiation was much less marked, presumably due to the greater dispersal of cpDNA via pollen.     

Survival and growth patterns of white spruce (Picea glauca [Moench] Voss) rangewide provenances and their implications for climate change adaptation

Intraspecific assisted migration (ISAM) through seed transfer during artificial forest regeneration has been suggested as an adaptation strategy to enhance forest resilience and productivity under future climate. In this study, we assessed the risks and benefits of ISAM in white spruce based on long‐term and multilocation, rangewide provenance test data. Our results indicate that the adaptive capacity and growth potential of white spruce varied considerably among 245 range‐wide provenances sampled across North America; however, the results revealed that local populations could be outperformed by nonlocal ones. Provenances originating from south‐central Ontario and southwestern Québec, Canada, close to the southern edge of the species' natural distribution, demonstrated superior growth in more northerly environments compared with local populations and performed much better than populations from western Canada and Alaska, United States. During the 19–28 years between planting and measurement, the southern provenances have not been more susceptible to freezing damage compared with local populations, indicating they have the potential to be used now for the reforestation of more northerly planting sites; based on changing temperature, these seed sources potentially could maintain or increase white spruce productivity at or above historical levels at northern sites. A universal response function (URF), which uses climatic variables to predict provenance performance across field trials, indicated a relatively weak relationship between provenance performance and the climate at provenance origin. Consequently, the URF from this study did not provide information useful to ISAM. The ecological and economic importance of conserving white spruce genetic resources in south‐central Ontario and southwestern Québec for use in ISAM is discussed.     

The recent population expansion of boarfish,Capros aper (Linnaeus, 1758): interactions of climate,growth and recruitment

The objectives of this study were to evaluate whether temperature changes in the Northeast Atlantic influence the growth and recruitment dynamics of boarfish, Capros aper. Two geographically separate areas were examined, ‘north’ at the northern distribution range west of Ireland and ‘south’ on the main fishing grounds south of Ireland. No significant differences in length‐at‐age were observed between the two areas. Interannual otolith growth patterns were similar between the two areas with distinct years of faster and slower growth. In the ‘north’, no significant relationship between adult growth and temperature was observed, while growth in the ‘south’ was positively related to temperature up to approximately 16°C growth rates were suppressed in the years with temperatures above that. Recruitment showed a positive correlation with adult growth the previous year for the Spanish recruitment index only, suggesting spatial connectivity between the Celtic Sea and the Bay of Biscay. The age distributions were similar in both areas and despite the boarfish's longevity of >30 years, are dominated by the age classes corresponding to the years with high recruitment, suggesting that increased recruitment is responsible for the observed stock expansion.     

Leopard tortoises in southern Africa have greater genetic diversity in the north than in the south (Testudinidae)

In contrast to mammals, little is known about the phylogeographic structuring of widely distributed African reptile species. With the present study, we contribute data for the leopard tortoise (Stigmochelys pardalis). It ranges from the Horn of Africa southward to South Africa and westwards to southern Angola. However, its natural occurrence is disputed for some southern regions. To clarify the situation, we used mtDNA sequences and 14 microsatellite loci from 204 individuals mainly from southern Africa. Our results retrieved five mitochondrial clades; one in the south and two in the north‐west and north‐east of southern Africa, respectively, plus two distributed further north. Using microsatellites, the southern clade matched with a well‐defined southern nuclear cluster, whilst the two northern clades from southern Africa corresponded to another nuclear cluster with three subclusters. One subcluster had a western and central distribution, another occurred mostly in the north‐east, and the third in a small eastern region (Maputaland), which forms part of a biodiversity hotspot. Genetic diversity was low in the south and high in the north of our study region, particularly in the north‐east. Our results refuted that translocations influenced the genetic structure of leopard tortoises substantially. We propose that Pleistocene climatic fluctuations caused leopard tortoises to retract to distinct refugia in southern and northern regions and ascribe the high genetic diversity in the north of southern Africa to genetic structuring caused by the survival in three refuges and subsequent admixture, whereas tortoises in the south seem to have survived in only one continuous coastal refuge.     

The fishery for black scabbardfish (Aphanopus carbo Lowe, 1839) in the Portuguese continental slope

The black scabbardfish is a deep-sea teleost species with high commercial interest in the NE Atlantic. While it is caught by large offshore trawlers in the North of Europe, in the south it is captured by artisanal longlines. On the Portuguese continental slope, in the south of ICES Division IXa, a longline fleet has been engaged in the capture of black scabbardfish for more than 20 years. This paper reviews and analyses the features of this fishery from a temporal perspective based on information from different data sources. The fishery is based on the port of Sesimbra, where a family type fishing community strives to keep catches profitable without altering the artisanal features that characterizes the fleet. Although the fleet has experienced some technical improvements through time, the fishing operations and strategy have changed very little. Fishing occurs at particular areas on the slope, where individual vessels have their own fishing grounds. Annual landings surpassed 4,000 tonnes in the early 1990’s, but decreased to nearly 3,000 tonnes and have remained at this level since 2000. The length structure of the landings was stable from 2000 to 2005, with a modal (total length) class of 1,090 mm. Monthly landings per unit effort estimated by a generalized linear model did not present any marked trend for the period 2000–2004. The landings reported in mainland Portugal were compared with the ones from other regions of the ICES area and the different characteristics of the fisheries operating in the north and south of Europe were analysed.     

Ecological niche modelling confirms potential north‐east range expansion of the nine‐banded armadillo (Dasypus novemcinctus) in the USA

The distribution of the nine banded armadillo (Dasypus novemcinctus), the only species in the family Dasypodidae found in the USA, has expanded greatly since the species was first recorded in southern Texas in 1849. Currently, the range of D. novemcinctus includes 15 states in the USA. Previous studies on the geographical expansion of this species, based on physiological experiments and distribution surveys, revealed a possible western moisture limit, a northern temperature limit, and potential north‐eastward range expansion in the USA. We applied an ecological niche modelling approach and produced a potential distribution map of D. novemcinctus with comparable western (102 °W) and northern (40 °N) limits, and confirmed the possibility of further north‐east range expansion to climatically suitable areas in the USA.     

Age, growth and mortality of red bandfish, Cepola macrophthalma (L.), in the western Aegean Sea (Greece)

Age, growth and mortality of red bandfish, Cepola macrophrhalma (L.), from the western Aegean Sea were studied using the otoliths from 1113 fish and the length and weight of 3351 fish. Two regions were established (north and south of the Euripos Strait). The red bandfish grows allometrically (slope of length-weight regressions ≤2) and relatively rapidly until age 4 or 5 years. It is proposed that this rapid linear growth represents an adaptation which evolved to reduce the vulnerability of red bandfish to whole-fish swallowers. Back-calculated lengths-at-age showed significant differences between regions and between sexes. Males were larger than females after age 2 years. Maximum age of northern fish was also higher than southern fish (8 and 5 years respectively). Southern fish grow at a faster rate (K=0.379) but to a significantly smaller size (L_x=424.8mm t.l. , W_x=25.1 g) than northern fish (K=0.214, L_x=676.1 mm t.l. , W_x=90.9 g). Mortality rates (total and natural instantaneous mortalities) of southern fish were also higher than northern fish. We suggest that these variations represent responses to the different conditions of temperature and food availability prevailing in the two regions.     

Reproduction and growth of Aphanopus carbo and A. intermedius (Teleostei: Trichiuridae) in the northeastern Atlantic

Commercial fishery landings of the black scabbardfish, Aphanopus carbo, from the Madeira mid‐water drifting longline fleet (northeastern Atlantic) were studied for 2 years within a Portuguese government programme for fishing management. The process noted that 20% of the total catch corresponded to the intermediate scabbardfish, Aphanopus intermedius. Length‐frequency distribution, and age, growth and reproduction of both species were analysed and compared. The results revealed significant differences in age and growth in influencing the length‐frequency distributions. Intermediate scabbardfish attained the largest size (148 cm TL) and age (15 years). However, the two species had similar reproductive strategies. According to data published on A. carbo in this region, it is thought to be very likely that previous studies unintentionally mixed both species together in their analyses.     

Comparing two fish sampling standards over time: largely congruent results but with caveats

相似文献   

Factors influencing the spatial distribution of zooplankton and fish in Loch Ness, UK

• 1 The vertical and horizontal distribution of phytoplankton, zooplankton and fish in Loch Ness, Scotland, were monitored during one day‐time and one night‐time survey in July 1992. The vertical samples were collected at a site located at the northern end of the loch and the horizontal samples along a longitudinal transect.
• 2 The vertical distribution surveys demonstrated that the phytoplankton, the zooplankton and the fish were concentrated in the top 30 m of water above the seasonal thermocline. Within this layer, Cyclops stayed much closer to the surface than Eudiaptomus but both species moved towards the surface at night.
• 3 The most important factor influencing the horizontal distribution of the phytoplankton was the north‐ south gradient in productivity. The sub‐catchments surrounding the north basin contain a greater proportion of arable land than those to the south and the concentrations of nitrate‐nitrogen and phytoplankton chlorophyll increased systematically from south to north.
• 4 Zooplankton distribution patterns were influenced by wind‐induced water movements and the dispersion of allochthonous material from the main inflows. The highest concentrations of Cyclops were recorded in the north, where there was more phytoplankton, and the highest concentrations of Eudiaptomus in the south, where there were higher concentrations of non‐algal particulates.
• 5 There was no spatial correlation between total zooplankton and total fish abundance but the highest concentrations of small (1–5 cm) fish were recorded in the south where there was a large patch of Eudiaptomus. The number of Eudiaptomus at specific locations within this patch were, however, negatively correlated with the numbers of small fish. These results suggest that the fish were actively foraging within the patch and were depleting their zooplankton prey in the areas where they were most abundant.

     

Is climate warming more consequential towards poles? The phenology of Lepidoptera in Finland

The magnitude and direction of phenological shifts from climate warming could be predictably variable across the planet depending upon the nature of physiological controls on phenology, the thermal sensitivity of the developmental processes and global patterns in the climate warming. We tested this with respect to the flight phenology of adult nocturnal moths (3.33 million captures of 334 species) that were sampled at sites in southern and northern Finland during 1993–2012 (with years 2005–2012 treated as an independent model validation data set). We compared eight competing models of physiological controls on flight phenology to each species and found strong support for thermal controls of phenology in 66% of the species generations. Among species with strong thermal control of phenology in both the south and north, the average development rate was higher in northern vs. southern populations at 10 °C, but about the same at 15 and 20 °C. With a 3 °C increase in temperature (approximating A2 scenario of IPPC for 2090–2099 relative to 1980–1999) these species were predicted to advance their phenology on average by 17 (SE ± 0.3) days in the south vs. 13 (±0.4) days in the north. The higher development rates at low temperatures of poleward populations makes them less sensitive to climate warming, which opposes the tendency for stronger phenological advances in the north from greater increases in temperature.     

The distribution of Athetis lepigone and prediction of its potential distribution based on GARP and MaxEnt

Athetis lepigone (Möschler) is a new agronomic pest which has caused serious damages to summer maize in China. In order to effectively monitor it, it is necessary to carry‐out a worldwide investigation on its potential geographical distribution. In this study, we give two ecological niche models, Genetic Algorithm for Rule‐set Production and Maximum Entropy (MaxEnt), to predict the potential geographical distribution of A. lepigone. The results indicate that the suitable areas for A. lepigone are mainly in China (Beijing, Tianjin, central and southern Hebei, northern Jiangsu, Shandong, most of Henan, northern Anhui, central and southern Shanxi, central and southern Shaanxi, small parts of north‐west in Hubei, eastern Gansu, parts of Ningxia, western of Xinjiang and Dandong, and Liaoning) and other Asian countries (South Korea, North Korea and Japan). Parts of Europe (south‐western Russia, Ukraine, Moldova, Romania, Hungary, Bulgaria, Slovenia, and Croatia, Bosnia and Herzegovina, Serbia, parts of Austria, western Poland, the Czech Republic, Germany, northern Italy, Denmark, western Lithuania, Latvia, Estonia, southern Finland and Sweden) are also highly suitable for A. lepigone. In addition, in the states of the USA including Michigan, New York, Chicago, Missouri, Ohio, Illinois and Indiana are also favourable for its occurrence although till now no reports about of this species has been recorded. A jackknife test in MaxEnt showed that the mean temperature of the driest quarter was the most important environmental variable affecting the distribution of this pest.     

Hook performance and selectivity of Eurasian perch,Perca fluviatilis (Linnaeus, 1758) in the Åland Archipelago,Finland

The effects of two hook‐types (circle or J hook) fished with two different baits (red worms Eisenia spp., or cyprinid fishes) were examined for deep‐hooking, catch rate, mean total length, and selectivity of Eurasian perch, Perca fluviatilis, in the Åland Archipelago, Finland. The perch population available to hook‐and‐line gear was sampled with multipanel gillnets, with panels ranging from 10 to 60 mm in stretched mesh diameter. The effect of each hook then was examined for each bait by conditioning the size distributions of hook‐and‐line catches on the size distribution of perch collected with gillnets to estimate selectivity. The proportion of deeply‐hooked perch was 4.8 times greater for J hooks (0.21) than circle hooks (0.04) baited with worms and 3.7 times greater for J hooks (0.33) than circle hooks (0.09) baited with fish. Catch rates were significantly different between the two baits but not between the two hooks, with the greatest catch rate observed for the circle hook baited with worms. There was no significant difference in perch total length between the two hooks, but there was a significant difference between the two baits, with the fish bait catching significantly larger perch. Perch selection peaks between the two hooks differed by only 13.8 mm, but the selection peak with fish bait (236.4 mm TL) was 119.9 mm greater than the selection peak with the worm bait (116.5 mm TL). A substantially lesser incidence of deep hooking support the conservation benefits of circle hook use for perch, especially when catch‐and‐release is practiced. Using fish baits similar in size to those used in this experiment can avoid potentially harmful interactions between relatively small perch and anglers who place little importance on catch motives but whose satisfaction is still catch‐dependent.     

Clinal adaptation and adaptive plasticity in Artemisia californica: implications for the response of a foundation species to predicted climate change

Local adaptation and plasticity pose significant obstacles to predicting plant responses to future climates. Although local adaptation and plasticity in plant functional traits have been documented for many species, less is known about population‐level variation in plasticity and whether such variation is driven by adaptation to environmental variation. We examined clinal variation in traits and performance – and plastic responses to environmental change – for the shrub Artemisia californica along a 700 km gradient characterized (from south to north) by a fourfold increase in precipitation and a 61% decrease in interannual precipitation variation. Plants cloned from five populations along this gradient were grown for 3 years in treatments approximating the precipitation regimes of the north and south range margins. Most traits varying among populations did so clinally; northern populations (vs. southern) had higher water‐use efficiencies and lower growth rates, C : N ratios and terpene concentrations. Notably, there was variation in plasticity for plant performance that was strongly correlated with source site interannual precipitation variability. The high‐precipitation treatment (vs. low) increased growth and flower production more for plants from southern populations (181% and 279%, respectively) than northern populations (47% and 20%, respectively). Overall, precipitation variability at population source sites predicted 86% and 99% of variation in plasticity in growth and flowering, respectively. These striking, clinal patterns in plant traits and plasticity are indicative of adaptation to both the mean and variability of environmental conditions. Furthermore, our analysis of long‐term coastal climate data in turn indicates an increase in interannual precipitation variation consistent with most global change models and, unexpectedly, this increased variation is especially pronounced at historically stable, northern sites. Our findings demonstrate the critical need to integrate fundamental evolutionary processes into global change models, as contemporary patterns of adaptation to environmental clines will mediate future plant responses to projected climate change.     

Distribution,abundance and biology of polar cod, <Emphasis Type="Italic">Boreogadus saida</Emphasis>, in Iceland–East Greenland waters

Recent warming has caused a northern extension in the distribution of many southern fish species in Icelandic waters. Polar cod (Boreogadus saida) around Iceland are near the southern limit of their distribution, but are poorly studied in the area. Therefore, material sampled during demersal fish surveys in March 1985–2013 and in pelagic 0-group surveys in Iceland–East Greenland waters in August–September 1974–2003 was used to investigate their distribution, abundance and biology. Demersal polar cod were most often caught on the outer shelf to the north-west and north of Iceland, but during years of widest distribution and highest abundance, they were caught farther to the east on the northern shelf. Pelagic 0-group polar cod were only caught sporadically and mainly confined to the waters off the north-west shelf of Iceland and the East Greenland shelf (southern Denmark Strait). In demersal hauls, the number of stations with polar cod decreased with increasing bottom temperature and polar cod were most widely distributed in the years 1989, 1994 and 1995. Highest numbers of demersal polar cod per haul were caught at temperatures of ?0.5 to 2.5 °C and at 200–450 m depth. The length of demersal polar cod ranged from 5 to 32 cm, while the fish caught in the pelagic trawl ranged from 2.2 to 19 cm. The polar cod in the subarctic waters north of Iceland most likely originate from the waters off East Greenland and further warming and decline in sea ice may eventually lead to the disappearance of polar cod from Icelandic waters.     

Monitoring demographics of a commercially exploited population of shovelnose sturgeon in the Wabash River,Illinois/Indiana,USA

Shovelnose sturgeon (Scaphirhynchus platorynchus, Rafinesque, 1820) in the Wabash River, Illinois/Indiana, USA, provide an important recreational sport and commercial caviar fishery. In fact, it is one of the last commercially viable populations for sturgeon roe harvest. Due to increased demand in the caviar trade and endangered species legislation that protect shovelnose sturgeon in only a portion of their range, efforts of the roe harvest market may continue to divert toward unprotected populations like the shovelnose sturgeon in the Wabash River. Previous studies have shown that increased harvest pressure in this species can affect the age‐at‐maturation and result in recruitment overfishing. Therefore, it is important to closely and continuously monitor commercially exploited populations. Over the past decade (2007–2016), 13,170 shovelnose sturgeon were sampled with boat electroshocking, hoop nets, drift nets, trotlines, and benthic electrified trawls. Captured fish ranged from 61 to 910 mm fork length (FL; mean = 668 mm), with very few fish less than 550 mm FL. Although fish were found to be in a healthy condition (mean relative weight = 87), there was a decrease in the mean condition over time. In addition, we saw declines in mean FL, weight of roe‐per‐fish, and size‐at‐maturity for female fish directly impacted by harvest. The decline of these population parameters, coupled with an increase in total annual mortality and a truncated age frequency distribution, suggest that harvest is negatively impacting the demographics and recruitment of shovelnose sturgeon in the Wabash River. Considering the downward trajectory of population dynamics and high estimates of mortality, their resiliency to continued harvest and environmental changes will be limited.