Life in 3-D: life history strategies in tunas, mackerels and bonitos

The scombrids (tunas, bonitos, Spanish mackerels and mackerels) sustain some of the most important fisheries in the world and their sustainable management depends on better understanding of their life history strategies. Here, we first assemble life history information on maximum size, growth, longevity, maturity, fecundity and spawning duration and interval for all scombrid species. Second we characterize their life history patterns and trait co-variation and evaluate how many principal axes of trait variation underlie scombrid life history strategies. Most of their life history variation can be explained along three axes or dimensions: size, speed, and reproductive schedule. Body size governs the first axis ranking species along a small-large continuum. The second axis was mostly influenced by time-related traits, such as longevity, growth rates, spawning duration, time between spawning events, ranking species along a slow-fast continuum of life histories. Scombrid species with the slowest life histories such as Atlantic bluefin tuna Thunnus thynnus and Atlantic mackerel Scomber scombrus tend to inhabit more temperate waters while species with faster life histories such as yellowfin tuna Thunnus albacares and short mackerel Rastrelliger brachysoma are typically found in more tropical waters. The third axis comprises the negative relationship between number of eggs produced at length of maturity and rate in gain of fecundity with size describing the schedule of reproductive allocation which reflects a fundamental trade-off between reproduction and growth. Finally, in addition we show that the life history strategies of scombrids conform more closely to the Periodic and Opportunistic strategists within the triangular model of fish life histories.     

Gill morphometrics in relation to gas transfer and ram ventilation in high‐energy demand teleosts: Scombrids and billfishes

This comparative study of the gill morphometrics in scombrids (tunas, bonitos, and mackerels) and billfishes (marlins, swordfish) examines features of gill design related to high rates of gas transfer and the high‐pressure branchial flow associated with fast, continuous swimming. Tunas have the largest relative gill surface areas of any fish group, and although the gill areas of non‐tuna scombrids and billfishes are smaller than those of tunas, they are also disproportionally larger than those of most other teleosts. The morphometric features contributing to the large gill surface areas of these high‐energy demand teleosts include: 1) a relative increase in the number and length of gill filaments that have, 2) a high lamellar frequency (i.e., the number of lamellae per length of filament), and 3) lamellae that are long and low in profile (height), which allows a greater number of filaments to be tightly packed into the branchial cavity. Augmentation of gill area through these morphometric changes represents a departure from the general mechanism of area enhancement utilized by most teleosts, which lengthen filaments and increase the size of the lamellae. The gill design of scombrids and billfishes reflects the combined requirements for ram ventilation and elevated energetic demands. The high lamellar frequencies and long lamellae increase branchial resistance to water flow which slows and streamlines the ram ventilatory stream. In general, scombrid and billfish gill surface areas correlate with metabolic requirements and this character may serve to predict the energetic demands of fish species for which direct measurement is not possible. The branching of the gill filaments documented for the swordfish in this study appears to increase its gill surface area above that of other billfishes and may allow it to penetrate oxygen‐poor waters at depth. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Structural adaptations for ram ventilation: Gill fusions in scombrids and billfishes

For ram‐gill ventilators such as tunas and mackerels (family Scombridae) and billfishes (families Istiophoridae, Xiphiidae), fusions binding the gill lamellae and filaments prevent gill deformation by a fast and continuous ventilatory stream. This study examines the gills from 28 scombrid and seven billfish species in order to determine how factors such as body size, swimming speed, and the degree of dependence upon ram ventilation influence the site of occurrence and type of fusions. In the family Scombridae there is a progressive increase in the reliance on ram ventilation that correlates with the elaboration of gill fusions. This ranges from mackerels (tribe Scombrini), which only utilize ram ventilation at fast cruising speeds and lack gill fusions, to tunas (tribe Thunnini) of the genus Thunnus, which are obligate ram ventilators and have two distinct fusion types (one binding the gill lamellae and a second connecting the gill filaments). The billfishes appear to have independently evolved gill fusions that rival those of tunas in terms of structural complexity. Examination of a wide range of body sizes for some scombrids and billfishes shows that gill fusions begin to develop at lengths as small as 2.0 cm fork length. In addition to securing the spatial configuration of the gill sieve, gill fusions also appear to increase branchial resistance to slow the high‐speed current produced by ram ventilation to distribute flow evenly and optimally to the respiratory exchange surfaces. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Fecundity of Atlantic sturgeon (Acipenser oxyrinchus Mitchill, 1815) captured by the commercial fishery in the Saint John River,New Brunswick,Canada

Knowledge of fecundity in fishes is important for understanding life history, modeling population dynamics, developing fisheries management and determining conservation status. Sturgeon species are known for their high fecundity but to date only two populations of Atlantic sturgeon (Acipenser oxyrinchus Mitchill, 1815) have been examined for this aspect of their biology. The species is protected in most regions of eastern North America but the Saint John River, New Brunswick supports one of the two remaining commercial fisheries in Canada. Sixty females selected by this fishery were sampled for total length, weight, age, gonad weight and egg number. Absolute fecundity ranged from 153,630 to 1,306,626 eggs, with a mean fecundity of 582,832 ± 261,806 eggs. Absolute fecundity exhibited a significant positive relationship to increasing body weight but not to total length or age. Relative fecundity ranged from 129 to 216 eggs/g gonad weight and although it declined with both increasing body size and age the relationships were not significant. The Gonadosomatic Index (GSI) ranged from 1.3% to 27.0% with a mean of 7.8% ± 4.94 SD. Relative fecundity found in this study was higher and mean GSI lower than reported for other Atlantic sturgeon populations and other sturgeon species. Differences were probably due to location of the fishery downstream from spawning sites and selection of pre-spawning females by the commercial fishery.     

Oligonucleotide indexing of DNA barcodes: identification of tuna and other scombrid species in food products

Background  

DNA barcodes are a global standard for species identification and have countless applications in the medical, forensic and alimentary fields, but few barcoding methods work efficiently in samples in which DNA is degraded, e.g. foods and archival specimens. This limits the choice of target regions harbouring a sufficient number of diagnostic polymorphisms. The method described here uses existing PCR and sequencing methodologies to detect mitochondrial DNA polymorphisms in complex matrices such as foods. The reported application allowed the discrimination among 17 fish species of the Scombridae family with high commercial interest such as mackerels, bonitos and tunas which are often present in processed seafood. The approach can be easily upgraded with the release of new genetic diversity information to increase the range of detected species.     

Unique adaptations of the metabolic biochemistry of tunas and billfishes for life in the pelagic environment

Synopsis Virtually all characteristics of tunas and billfishes reflect their highly charged lifestyles as apex predators in the oceanic pelagic environment. The adaptations they possess for efficient and rapid swimming, efficient and rapid food processing, turnover of nutrients and storage and mobilization of internal fuel supplies, and for rapid recovery rates, are discussed. Overall, tunas and billfishes are designed for high performance, at both sustainable and burst swimming speeds, but there are several differences between tunas and billfishes. Tunas' aerobic metabolic capacities exceed those of ectothermic fishes, including billfishes and other scombrids, by virtue of their elevated red muscle temperatures, and because heart and white muscle aerobic capacities are significantly greater in tunas. The adaptations for high performance involve some costs, including the need for a constant high energy input to sustain high metabolic rates, high activity levels, and endothermy, Yet, tunas and billfishes have adopted successful lifestyles, as evidenced by their large numbers and biomass within the marine environment. Although our knowledge of these fishes has increased dramatically during the past 15 years, there are major gaps in our understanding of the metabolic biochemistry and physiology of these fishes, and these are highlighted so that additional research can be directed towards filling these gaps.Paper from the International Union of Biological Societies symposium The biology of tunas and billfishes: an examination of life on the knife edge, organized by Richard W. Brill and Kim N. Holland.     

A review of the biology,ecology, behavior and conservation status of the dusky grouper, <Emphasis Type="Italic">Epinephelus marginatus</Emphasis> (Lowe 1834)

The dusky grouper is a large-bodied marine species usually associated with rocky substrates and reefs currently experiencing increasing anthropogenic pressures. Here, we critically evaluate studies conducted during the last few decades on this endangered species, focusing on its biology, ecology, behavior, stock structure and fisheries, and identify future research directions to fill current knowledge gaps. This species plays an important functional role in the rocky bottom habitats in which it lives, usually occupying the highest trophic levels and with the potential role as a keystone species in some ecosystems. It has a complex life cycle characterized by a high longevity, a slow growth rate and a monandric protogynous hermaphroditic mode of reproduction. The species also exhibits spawning aggregation behavior, which makes it more vulnerable to fishing activity. Aside from its ecological importance, this species has high commercial value for professional fishers and is also highly prized by recreational spearfishers. These biological features, associated with increasing fishing pressures throughout most of its geographical distribution, have led to Epinephelus marginatus being included since 2004 in the IUCN Red List of Threatened Species as Endangered. Despite a large amount of research being conducted on this species over the last five decades or so, using a number of different tools and techniques, key questions regarding its biology still remain unanswered. In terms of conservation and management plans designed to safeguard this endangered species, we identified the crucial need to improve fisheries landing statistics for the species and to increase the number of marine protected areas within its distribution beyond the Mediterranean Sea.     

Reproduction and larval biology in tunas,and the importance of restricted area spawning grounds

Tunas show a wide variety of life history strategies, spatial distributions and migratory behaviors, yet they share a common trait of spawning only in tropical and sub-tropical regions. The warm-water tuna species generally show significant overlap between spawning and feeding grounds, and longer spawning seasons of several months to near year-round. In contrast, the cool-water bluefin tunas migrate long distances between feeding and spawning grounds, and may spawn over periods as short as 2 months. Here, we examine the spatial distributions of tuna larvae in the world’s oceans, and examine interspecific differences in the light of adult behaviors and larval ecology. We discuss the links between larval tuna and their oceanographic environments and relate these to current knowledge of larval growth, feeding and trophodynamics, with a focus on the better-studied bluefin tunas. We show that larval tunas have moderate to fast growth rates and selective feeding habits, and thus appear to be adapted for survival in warm, oligotrophic seas. We also examine the challenges of sustainably managing species which migrate across multiple management boundaries to reach spatio-temporally restricted spawning grounds and discuss the previous and future anthropogenic impacts on tuna spawning areas.     

Evolution and consequences of endothermy in fishes

Regional endothermy, the conservation of metabolic heat by vascular countercurrent heat exchangers to elevate the temperature of the slow-twitch locomotor muscle, eyes and brain, or viscera, has evolved independently among several fish lineages, including lamnid sharks, billfishes, and tunas. All are large, active, pelagic species with high energy demands that undertake long-distance migrations and move vertically within the water column, thereby encountering a range of water temperatures. After summarizing the occurrence of endothermy among fishes, the evidence for two hypothesized advantages of endothermy in fishes, thermal niche expansion and enhancement of aerobic swimming performance, is analyzed using phylogenetic comparisons between endothermic fishes and their ectothermic relatives. Thermal niche expansion is supported by mapping endothermic characters onto phylogenies and by combining information about the thermal niche of extant species, the fossil record, and paleoceanographic conditions during the time that endothermic fishes radiated. However, it is difficult to show that endothermy was required for niche expansion, and adaptations other than endothermy are necessary for repeated diving below the thermocline. Although the convergent evolution of the ability to elevate slow-twitch, oxidative locomotor muscle temperatures suggests a selective advantage for that trait, comparisons of tunas and their ectothermic sister species (mackerels and bonitos) provide no direct support of the hypothesis that endothermy results in increased aerobic swimming speeds, slow-oxidative muscle power, or energetic efficiency. Endothermy is associated with higher standard metabolic rates, which may result from high aerobic capacities required by these high-performance fishes to conduct many aerobic activities simultaneously. A high standard metabolic rate indicates that the benefits of endothermy may be offset by significant energetic costs.     

The evolution of thunniform locomotion and heat conservation in scombrid fishes: New insights based on the morphology of Allothunnus fallai

Two significant functional differences–a more anterior and internally positioned red myo‐tomal muscle mass and modification of the red‐muscle vascular supply to form counter‐current heat exchangers–distinguish the tunas (tribe Thunnini) from other species in the teleost family Scombridae. Neither of these characteristics is found in the bonitos (tribe Sardini), the sister group to the Thunnini. The most recent scombrid classification places the slender tuna, Allothunnus fallai, in the tribe Sardini, but some earlier studies suggested that this species should be a member of the Thunnini. Allothunnus fallai does not possess the lateral subcutaneous arteries and veins or the lateral heat‐exchanging retia typical of tunas. However, we have found that this species has a highly modified central circulation (dorsal aorta, post cardinal vein, and associated branch vessels) similar to the central heat‐exchanging retia of certain tunas, an enlarged haemal arch to accommodate this structure, and the anterior, internal placement of red muscle characteristic of tunas. With these new characters, phylogenetic reconstructions based on parsimony place A. fallai as the sister taxon to the tunas, establish that it is the most basal tuna species, and support the hypothesis that the derivation of tunas from a bonito‐like ancestor occurred through selection for an integrated set of characteristics affecting locomotion and endothermy. The major features of this hypothesis are as follows. (1) Selection for continuous, steady, and efficient swimming resulted in changes in body shape (the result of enlargement of the anterior myotomes, the anterior and internal shift of red muscle, and a narrowing of the caudal peduncle) which increased streamlining and led to the adoption of the thunniform swimming mode unique to the tunas. (2) Alterations in blood supply necessitated by the anterior shift in red muscle led to the interdigitation of numerous arterial and venous branches which set the stage for heat conservation. (3) The evolution of endothermy, together with thunniform swimming, contributed significantly to the ecological radiation and diversification of tunas during the Early Tertiary Period. Our studies of A fall thus suggest that the shift in red muscle position and changes in central circulation preerded the evolution of red‐muscle endothermy. Co‐evolutionary changes in red muscle quantity and distribution and in vascular specializations for heat conservation have led to different macroevolutionary trajectories among the now five genera and 14 tuna species of tunas and appear to reflect the influence of changing paleocological and paleoccanographic conditions, including cooling, that occurred in the Tertiary     

Longtail tuna,Thunnus tonggol (Bleeker, 1851): a global review of population dynamics,ecology, fisheries,and considerations for future conservation and management

Longtail tuna (Thunnus tonggol) is a neritic species that supports commercial, artisanal and recreational fisheries throughout the Indo-Pacific region. Historically receiving little attention by commercial fisheries, the global annual catch of longtail tuna has steadily risen from around 30,000 t in the early 1980s to exceeding 200,000 t since 2004, reaching a peak of 291,264 t in 2007, and was 281,613 t in 2017. Catches of longtail tuna in the Indian Ocean now exceed catches of principal commercial target species, such as albacore and bigeye tunas. A sequence of stock assessments undertaken throughout the species’ range since the late 1980s persistently indicated that at least three of the four stocks defined in this paper are likely to have been, and most likely are currently, subject to overfishing and overfished as a result of excess fishing effort on this relatively slow-growing and long-lived tuna species. As the spawning biomass of principal tuna target species continue to decline in both the Indian and western and central Pacific Oceans, the increasing catches of longtail tuna, other neritic tunas, and seerfishes is worrisome. Few conservation and management measures (CMMs) are currently in place specifically for longtail tuna, although in recent years some coastal States, Regional Fishery Bodies, and tuna Regional Fisheries Management Organisations have begun to develop initiatives to improve the catch and biological data quality for longtail tuna and sympatric species of neritic tunas and tuna-like species. This paper provides a global review of biological, ecological and fishery information to provide researchers, fishery managers and policy makers with the most current information from which to begin to guide future stock assessment and the development of CMMs for longtail tuna.

     

Life history and population size variability in a relict plant. Different routes towards long-term persistence

A central tenet of conservation biology is that population size affects the persistence of populations. However, many narrow endemic species combine small population ranges and sizes with long persistence, thereby challenging this tenet. I examined the performance of three different-sized populations of Petrocoptis pseudoviscosa (Caryophyllaceae), a palaeoendemic rupicolous herb distributed along a small valley in the Spanish Pyrenees. Reproductive and demographic parameters were recorded over 6 years, and deterministic and stochastic matrix models were constructed to explore population dynamics and extinction risk. Populations differed greatly in structure, fecundity, recruitment, survival rate, and life span. Strong differentiation in life-history parameters and their temporal variability resulted in differential population vulnerability under current conditions and simulated global changes such as habitat fragmentation or higher climatic fluctuations. This study provides insights into the capacity of narrow endemics to survive both at extreme environmental conditions and at small population sizes. When dealing with species conservation, the population size–extinction risk relationship may be too simplistic for ancient, ecologically restricted organisms, and some knowledge of life history may be most important to assess their future.     

Gametogenesis and fecundity of <Emphasis Type="Italic">Acropora tenella</Emphasis> (Brook 1892) in a mesophotic coral ecosystem in Okinawa,Japan

Mesophotic coral ecosystems (below 30–40 m depth) host a large diversity of zooxanthellate coral communities and may play an important role in the ecology and conservation of coral reefs. Investigating the reproductive biology of mesophotic corals is important to understand their life history traits. Despite an increase in research on mesophotic corals in the last decade, their reproductive biology is still poorly understood. Here, gametogenesis and fecundity of the Indo-Pacific mesophotic coral, Acropora tenella, were examined in an upper mesophotic reef (40 m depth) in Okinawa, Japan for the first time. Acropora tenella is a hermaphrodite with a single annual gametogenic cycle, and both oogenesis and spermatogenesis occurring for 11–12 and 5–6 months, respectively. Timing of spawning of this species was similar to other shallow Acropora spp. in the region. However, colonies had longer gametogenic cycles and less synchronous gamete maturation compared to shallow acroporids with spawning extended over consecutive months. Both the polyp fecundity (number of eggs per polyp) and gonad index (defined as the number of eggs per square centimeter) of A. tenella were lower than most acroporids. Our findings contribute to understanding of the life history of corals on mesophotic reefs and suggest that the reproductive biology of upper mesophotic corals is similar to that of shallow-water corals.     

Identification of small juvenile scombrids from northwest tropical Australia using mitochondrial DNA cytochrome <Emphasis Type="Italic">b</Emphasis> sequences

Small juveniles of the nine species of scombrids in Australian waters are morphologically similar to one another and, consequently, difficult to identify to species level. We show that the sequence of the mitochondrial DNA cytochrome b gene region is a powerful tool for identification of these young fish. Using this method, we identified 50 juvenile scombrids collected from Exmouth Bay, Western Australia. Six species of scombrids were apparent in this sample of fish: narrow-barred Spanish mackerel (Scomberomorus commerson), Indian mackerel (Rastrelliger kanagurta), frigate tuna (Auxis thazard), bullet tuna (Auxis rochei), leaping bonito (Cybiosarda elegans), and kawakawa (Euthynnus affinis). The presence of Indian mackerel, frigate tuna, leaping bonito, and kawakawa is the first indication that coastal waters may be an important spawning habitat for these species, although offshore spawning may also occur. The occurrence of small juvenile S. commerson was predicted from the known spawning patterns of that species, but other mackerel species (Scomberomorus munroi, Scomberomorus queenslandicus, Scomberomorus semifasiciatus) likely to be spawning during the sampling period were not detected among the 50 small juveniles analyzed here.     

A New Genus and Species of Scombrid Fish (Perciformes,Scombroidei, Scombridae) from the Lower Oligocene of the Caucasus

A new genus and species of scombrid fish Caucombrus histiopterygius gen. et sp. nov. is described from the Lower Oligocene (Lower Maikopian, Pshekha regional state) of the north-western Caucasus (Russia) and Abkhazia (rivers Belaya, Pshekha and Gumista). The new taxon differs from the other known Scombridae, both fossil and Recent, by the high and sail-like first dorsal fin. Osteological characters of Caucombrus gen. nov. indicate its relationship with the Spanish mackerels (subfamily Scomberomorinae). It is assumed that either C. histiopterygius gen. et sp. nov. or the related species of Caucombrus gen. nov. inhabited the Mid-Oligocene sea basin in the territory of the Rhine Graben (Germany).     

Phylogenetic effects on functional traits and life history strategies of Australian freshwater fish

Understanding the biogeographic and phylogenetic basis to interspecific differences in species’ functional traits is a central goal of evolutionary biology and community ecology. We quantify the extent of phylogenetic influence on functional traits and life‐history strategies of Australian freshwater fish to highlight intercontinental differences as a result of Australia's unique biogeographic and evolutionary history. We assembled data on life history, morphological and ecological traits from published sources for 194 Australian freshwater species. Interspecific variation among species could be described by a specialist–generalist gradient of variation in life‐history strategies associated with spawning frequency, fecundity and spawning migration. In general, Australian fish showed an affinity for life‐history strategies that maximise fitness in hydrologically unpredictable environments. We also observed differences in trait lability between and within life history, morphological and ecological traits where in general morphological and ecological traits were more labile. Our results showed that life‐history strategies are relatively evolutionarily labile and species have potentially evolved or colonised in freshwaters frequently and independently allowing them to maximise population performance in a range of environments. In addition, reproductive guild membership showed strong phylogenetic constraint indicating that evolutionary history is an important component influencing the range and distribution of reproductive strategies in extant species assemblages. For Australian freshwater fish, biogeographic and phylogenetic history contribute to broad taxonomic differences in species functional traits, while finer scale ecological processes contribute to interspecific differences in smaller taxonomic units. These results suggest that the lability or phylogenetic relatedness of different functional traits affects their suitability for testing hypothesis surrounding community level responses to environmental change.     

First estimates of metabolic rate in Atlantic bluefin tuna larvae

Atlantic bluefin tuna is an iconic scombrid species with a high commercial and ecological value. Despite their importance, many physiological aspects, especially during the larval stages, are still unknown. Metabolic rates are one of the understudied aspects in scombrid larvae, likely due to challenges associated to larval handling before and during respirometry trials. Gaining reliable estimates of metabolic rates is essential to understand how larvae balance their high growth needs and activity and other physiological functions, which can be very useful for fisheries ecology and aquaculture. This is the first study to (a) estimate the relationship between routine metabolic rate (RMR) and larval dry weight (DW) (mass scaling exponent) at a constant temperature of 26°C, (b) measure the RMR under light and darkness and (c) test whether the interindividual differences in the RMR are related to larval nutritional status (RNA/DNA and DNA/DW). The RMR scaled nearly isometrically with body size (b = 0.99, 0.60–31.56 mg DW) in contrast to the allometric relationship observed in most fish larvae (average b = 0.87). The results show no significant differences in larval RMR under light and darkness, suggesting similar larval activity levels in both conditions. The size explained most of the variability in RMR (97%), and nutritional condition was unrelated to the interindividual differences in routine metabolism. This is the first study to report the metabolic rates of Atlantic bluefin tuna larvae and discuss the challenges of performing bioenergetic studies with early life stages of scombrids.     

Marine Telemetry and the Conservation and Management of Risk to Seal Species in Canada and Australia

Marine telemetry expands the knowledge of the biology of marine species at risk: their life cycles, activities, interactions, habitats, and threats. Four seal species in Canada and Australia are faced with distinctive and divergent management problems. This article examines their conservation status, legal protection, and the role that telemetry has played, or could play, in providing previously unavailable information to help meet conservation goals. The value of telemetry data to minimize fisheries mortality of one species has been demonstrated in Australia. Despite there being significant telemetry data for the other species, policy and management have not yet responded.     

Phylogeny,extinction and conservation: embracing uncertainties in a time of urgency

Evolutionary studies have played a fundamental role in our understanding of life, but until recently, they had only a relatively modest involvement in addressing conservation issues. The main goal of the present discussion meeting issue is to offer a platform to present the available methods allowing the integration of phylogenetic and extinction risk data in conservation planning. Here, we identify the main knowledge gaps in biodiversity science, which include incomplete sampling, reconstruction biases in phylogenetic analyses, partly known species distribution ranges, and the difficulty in producing conservation assessments for all known species, not to mention that much of the effective biological diversity remains to be discovered. Given the impact that human activities have on biodiversity and the urgency with which we need to address these issues, imperfect assumptions need to be sanctioned and surrogates used in the race to salvage as much as possible of our natural and evolutionary heritage. We discuss some aspects of the uncertainties found in biodiversity science, such as the ideal surrogates for biodiversity, the gaps in our knowledge and the numerous available phylogenetic diversity-based methods. We also introduce a series of cases studies that demonstrate how evolutionary biology can effectively contribute to biodiversity conservation science.