Nonlethally assessing elasmobranch ontogenetic shifts in energetics.

Body condition is an important proxy for the overall health and energetic status of fishes. The classically used Fulton's condition factor requires length and mass measurements, but mass can be difficult to obtain in large species. Girth measurements can replace mass for wild pelagic sharks. However, girth-calculated condition has not been validated against Fulton's condition factor intraspecifically, across ontogeny or reproduction, or in a controlled setting. We used the epaulette shark (Hemiscyllium ocellatum), because they are amenable to captive reproduction, to track fine-scale body condition changes across life stages, oviparous reproduction and between condition indices. We measured four girths, total length and mass of 16 captive epaulette sharks across 1 year and tracked female reproduction daily. We also collected length and mass data from an additional 72 wild-caught sharks and 155 sharks from five previous studies and two public aquaria to examine the relationship between length and mass for this species. Even though data were derived from a variety of sources, a predictable length-mass relationship (R2 = 0.990) was achievable, indicating that combining data from a variety of sources could help overcome knowledge gaps regarding basic life history characteristics. We also found that condition factor decreased during early life stages, then increased again into adulthood, with predictable changes across the female reproductive cycle. Finally, we determined that both Fulton's and girth condition analyses were comparable. Outcomes from this study uniquely provide body condition changes across the complete life history, including fine-scale female reproductive stages, and validate the use of girths as a nonlethal whole-organism energetic assessment for fishes.

The upper thermal limit of epaulette sharks (Hemiscyllium ocellatum) is conserved across three life history stages, sex and body size.

Owing to climate change, most notably the increasing frequency of marine heatwaves and long-term ocean warming, better elucidating the upper thermal limits of marine fishes is important for predicting the future of species and populations. The critical thermal maximum (CTmax), or the highest temperature a species can tolerate, is a physiological metric that is used to establish upper thermal limits. Among marine organisms, this metric is commonly assessed in bony fishes but less so in other taxonomic groups, such as elasmobranchs (subclass of sharks, rays and skates), where only thermal acclimation effects on CTmax have been assessed. Herein, we tested whether three life history stages, sex and body size affected CTmax in a tropical elasmobranch, the epaulette shark (Hemiscyllium ocellatum), collected from the reef flats surrounding Heron Island, Australia. Overall, we found no difference in CTmax between life history stages, sexes or across a range of body sizes. Findings from this research suggest that the energetically costly processes (i.e. growth, maturation and reproduction) associated with the life history stages occupying these tropical reef flats do not change overall acute thermal tolerance. However, it is important to note that neither embryos developing in ovo, neonates, nor females actively encapsulating egg cases were observed in or collected from the reef flats. Overall, our findings provide the first evidence in an elasmobranch that upper thermal tolerance is not impacted by life history stage or size. This information will help to improve our understanding of how anthropogenic climate change may (or may not) disproportionally affect particular life stages and, as such, where additional conservation and management actions may be required.

Diel Rhythm and Thermal Independence of Metabolic Rate in a Benthic Shark.

Biological rhythms that are mediated by exogenous factors, such as light and temperature, drive the physiology of organisms and affect processes ranging from cellular to population levels. For elasmobranchs (i.e. sharks, rays, and skates), studies documenting diel activity and movement patterns indicate that many species are crepuscular or nocturnal in nature. However, few studies have investigated the rhythmicity of elasmobranch physiology to understand the mechanisms underpinning these distinct patterns. Here, we assess diel patterns of metabolic rates in a small meso-predator, the epaulette shark (Hemiscyllium ocellatum), across ecologically relevant temperatures and upon acutely removing photoperiod cues. This species possibly demonstrates behavioral sleep during daytime hours, which is supported herein by low metabolic rates during the day and a 1.7-fold increase in metabolic rates at night. From spring to summer seasons, where average average water temperature temperatures for this species range 24.5 to 28.5 °C, time of day, and not temperature, had the strongest influence on metabolic rate. These results indicate that this species, and perhaps other similar species from tropical and coastal environments, may have physiological mechanisms in place to maintain metabolic rate on a seasonal time scale regardless of temperature fluctuations that are relevant to their native habitats.

One hundred research questions in conservation physiology for generating actionable evidence to inform conservation policy and practice.

Environmental change and biodiversity loss are but two of the complex challenges facing conservation practitioners and policy makers. Relevant and robust scientific knowledge is critical for providing decision-makers with the actionable evidence needed to inform conservation decisions. In the Anthropocene, science that leads to meaningful improvements in biodiversity conservation, restoration and management is desperately needed. Conservation Physiology has emerged as a discipline that is well-positioned to identify the mechanisms underpinning population declines, predict responses to environmental change and test different in situ and ex situ conservation interventions for diverse taxa and ecosystems. Here we present a consensus list of 10 priority research themes. Within each theme we identify specific research questions (100 in total), answers to which will address conservation problems and should improve the management of biological resources. The themes frame a set of research questions related to the following: (i) adaptation and phenotypic plasticity; (ii) human-induced environmental change; (iii) human-wildlife interactions; (iv) invasive species; (v) methods, biomarkers and monitoring; (vi) policy, engagement and communication; (vii) pollution; (viii) restoration actions; (ix) threatened species; and (x) urban systems. The themes and questions will hopefully guide and inspire researchers while also helping to demonstrate to practitioners and policy makers the many ways in which physiology can help to support their decisions.

Future thermal regimes for epaulette sharks (Hemiscyllium ocellatum): growth and metabolic performance cease to be optimal.

Climate change is affecting thermal regimes globally, and organisms relying on their environment to regulate biological processes face unknown consequences. In ectotherms, temperature affects development rates, body condition, and performance. Embryonic stages may be the most vulnerable life history stages, especially for oviparous species already living at the warm edge of their distribution, as embryos cannot relocate during this developmental window. We reared 27 epaulette shark (Hemiscyllium ocellatum) embryos under average summer conditions (27 °C) or temperatures predicted for the middle and end of the twenty-first century with climate change (i.e., 29 and 31 °C) and tracked growth, development, and metabolic costs both in ovo and upon hatch. Rearing sharks at 31 °C impacted embryonic growth, yolk consumption, and metabolic rates. Upon hatch, 31 °C-reared sharks weighed significantly less than their 27 °C-reared counterparts and exhibited reduced metabolic performance. Many important growth and development traits in this species may peak after 27 °C and start to become negatively impacted nearing 31 °C. We hypothesize that 31 °C approximates the pejus temperature (i.e., temperatures at which performance of a trait begin to decline) for this species, which is alarming, given that this temperature range is well within ocean warming scenarios predicted for this species' distribution over the next century.

Reframing conservation physiology to be more inclusive, integrative, relevant and forward-looking: reflections and a horizon scan.

Applying physiological tools, knowledge and concepts to understand conservation problems (i.e. conservation physiology) has become commonplace and confers an ability to understand mechanistic processes, develop predictive models and identify cause-and-effect relationships. Conservation physiology is making contributions to conservation solutions; the number of 'success stories' is growing, but there remain unexplored opportunities for which conservation physiology shows immense promise and has the potential to contribute to major advances in protecting and restoring biodiversity. Here, we consider how conservation physiology has evolved with a focus on reframing the discipline to be more inclusive and integrative. Using a 'horizon scan', we further explore ways in which conservation physiology can be more relevant to pressing conservation issues of today (e.g. addressing the Sustainable Development Goals; delivering science to support the UN Decade on Ecosystem Restoration), as well as more forward-looking to inform emerging issues and policies for tomorrow. Our horizon scan provides evidence that, as the discipline of conservation physiology continues to mature, it provides a wealth of opportunities to promote integration, inclusivity and forward-thinking goals that contribute to achieving conservation gains. To advance environmental management and ecosystem restoration, we need to ensure that the underlying science (such as that generated by conservation physiology) is relevant with accompanying messaging that is straightforward and accessible to end users.

Wound healing in an elasmobranch fish is not impaired by high-CO2 exposure.

The purpose of this study was to test the effects of high CO2 exposure on wound healing rates in an elasmobranch fish (Urobatis jamaicensis). Small dermal injuries (8 mm biopsy) closed by 22 days post wounding with a decrease in haematocrit. High CO2 exposure (ΔpH = 1.4) did not influence healing rate or haematocrit. Combined, these data provide evidence that minimally invasive scientific procedures have short-term impacts on elasmobranch fishes even during exposure to a chronic stressor. Therefore, wound healing rates may not be strongly impacted by ocean acidification (ΔpH = 0.4).

Cortisol does not increase risk of mortality to predation in juvenile bluegill sunfish: A manipulative experimental field study.

The hypothalamic-pituitary-interrenal (HPI) or stress axis in teleost fishes produces their primary glucocorticoid, cortisol. Although generally an adaptive response, prolonged HPI axis stimulation can impair organismal performance. Previous work has shown that stressed teleosts have higher mortality to predation than unstressed conspecifics, suggesting a role for HPI axis in modulating predator-prey interactions. Our current study investigated whether elevated cortisol levels altered the predation rate of a wild teleost fish, the bluegill sunfish (Lepomis macrochirus). Wild juvenile bluegill were given intraperitoneal implants of cocoa butter (i.e., sham), or cocoa butter containing cortisol or cortisol and the glucocorticoid receptor antagonist RU486. After 24 hr, fish were tethered along the bottom of the lake and their survival under natural predation was recorded following 24 hr. A subset of fish was used to validate the efficacy of cortisol implants in this setting. No treatment effect on survival was observed, suggesting that elevated cortisol has minimal involvement in mediating predator-prey interactions in this context. However, experimental fish may have demonstrated resiliency to physiological perturbations owing to the relatively acute duration of our experimental series, and negative effects might be manifested over a more chronic period.

Exercise intensity while hooked is associated with physiological status of longline-captured sharks.

Some shark populations face declines owing to targeted capture and by-catch in longline fisheries. Exercise intensity during longline capture and physiological status may be associated, which could inform management strategies aimed at reducing the impacts of longline capture on sharks. The purpose of this study was to characterize relationships between exercise intensity and physiological status of longline-captured nurse sharks (Ginglymostoma cirratum) and Caribbean reef sharks (Carcharhinus perezi). Exercise intensity of longline-captured sharks was quantified with digital cameras and accelerometers, which was paired with blood-based physiological metrics from samples obtained immediately post-capture. Exercise intensity was associated with physiological status following longline capture. For nurse sharks, blood pH increased with capture duration and the proportion of time exhibiting low-intensity exercise. Nurse sharks also had higher blood glucose and plasma potassium concentrations at higher sea surface temperatures. Associations between exercise intensity and physiological status for Caribbean reef sharks were equivocal; capture duration had a positive relation with blood lactate concentrations and a negative relationship with plasma chloride concentrations. Because Caribbean reef sharks did not appear able to influence blood pH through exercise intensity, this species was considered more vulnerable to physiological impairment. While both species appear quite resilient to longline capture, it remains to be determined if exercise intensity during capture is a useful tool for predicting mortality or tertiary sub-lethal consequences. Fisheries management should consider exercise during capture for sharks when developing techniques to avoid by-catch or reduce physiological stress associated with capture.

The effect of capture and handling stress in Lophius americanus in the scallop dredge fishery.

Capture and handling stress studies are considered a primary research priority, particularly for species and fisheries where discard rates are high, and/or for overfished stocks and species of concern. Lophius americanus, a commercially valuable finfish in New England, constitutes the second highest bycatch species within the sea scallop dredge fishery. Despite its commercial importance, no data exists on the capture and handling stress of monkfish for any gear type. Given these shortcomings, our goals were to evaluate the stress response of monkfish captured in scallop dredge gear by evaluating physical, behavioural and physiological responses to scallop fishing practices. While 80% of monkfish displayed little to no physical trauma, behavioural and physiological assessment indicated high levels of stress, especially as air exposure and tow duration increased. This finding suggests that the manifestation of stress in monkfish may be a cryptic response necessitating further research in addition to estimates of post-release mortality rates to appropriately advise fisheries management regarding the mortality of monkfish bycatch in the sea scallop fishery.

Influence of supraphysiological cortisol manipulation on predator avoidance behaviors and physiological responses to a predation threat in a wild marine teleost fish.

The stress axis in teleost fish attempts to maintain internal homeostasis in the face of allostatic loading. However, stress axis induction has been associated with a higher predation rate in fish. To date, the physiological and behavioral factors associated with this outcome are poorly understood. The purpose of the present study was to investigate the impact of experimental cortisol elevation on anti-predator behavior and physiological responses to predator presence. We hypothesized that semi-chronic cortisol elevation would increase susceptibility to predation by increasing stress-induced risk-taking behaviors. To test this hypothesis, schoolmaster snapper were given cocoa butter implants without cortisol (sham) or with cortisol (50 mg/kg body weight) and tethered to cover. Fish were exposed to either a lemon shark or control conditions for 15-min. Space use and activity were recorded throughout and fish were terminally sampled for blood. Cortisol implantation, relative to shams, resulted in higher blood glucose and plasma cortisol concentrations with a lower plasma lactate concentration. Shark exposure, relative to controls, elicited higher blood glucose and lactate concentrations but had no effect on plasma cortisol concentration. No interactions were detected between shark exposure and cortisol treatment for any physiological trait. Behavioral metrics, including shelter use and activity, were unaffected by either cortisol implantation or shark exposure. Physiological responses to cortisol implantation likely resulted from enhanced gluconeogenic activity, whereas alterations under predator exposure may have been the product of catecholamine mobilization. Further work should address context-specific influences of stress in mediating behavioral responses to predation.

Validation of a portable, waterproof blood pH analyser for elasmobranchs.

Quantifying changes in blood chemistry in elasmobranchs can provide insights into the physiological insults caused by anthropogenic stress, and can ultimately inform conservation and management strategies. Current methods for analysing elasmobranch blood chemistry in the field are often costly and logistically challenging. We compared blood pH values measured using a portable, waterproof pH meter (Hanna Instruments HI 99161) with blood pH values measured by an i-STAT system (CG4+ cartridges), which was previously validated for teleost and elasmobranch fishes, to gauge the accuracy of the pH meter in determining whole blood pH for the Cuban dogfish (Squalus cubensis) and lemon shark (Negaprion brevirostris). There was a significant linear relationship between values derived via the pH meter and the i-STAT for both species across a wide range of pH values and temperatures (Cuban dogfish: 6.8-7.1 pH 24-30°C; lemon sharks: 7.0-7.45 pH 25-31°C). The relative error in the pH meter's measurements was ~±2.7%. Using this device with appropriate correction factors and consideration of calibration temperatures can result in both a rapid and accurate assessment of whole blood pH, at least for the two elasmobranch species examined here. Additional species should be examined in the future across a wide range of temperatures to determine whether correction factors are universal.

An experimental evaluation of the role of the stress axis in mediating predator-prey interactions in wild marine fish.

The hypothalamic-pituitary-interrenal (HPI) axis, through corticosteroid secretion, is an integral mechanism regulating internal homeostasis when vertebrates are faced with a stressor. However, continued HPI-axis stimulation can produce homeostatic overload, where corticosteroids are detrimental to organismal function. This overload condition may play an important role in mediating predator-prey interactions, because chronically/previously stressed animals may have higher rates of predator-induced mortality. However, the mechanism(s) underlying this observation are unknown. Using fish as models, we hypothesized that chronic stress would increase predation susceptibility owing to a poor physiological state (e.g. homeostatic overload) with corresponding sub-optimal changes in predator-avoidance behaviour. As cortisol is also required in low quantities to help regulate basic metabolic functions in fish, we expected that a glucocorticoid receptor antagonist (GR; e.g. homeostatic failure) may produce similar effects. Schoolmaster snapper (Lutjanus apodus) were given intraperitoneal implants of cocoa butter impregnated with nothing (sham; 5ml/kg body weight (BW)), cortisol (50mg/kg BW) or the GR antagonist RU486 (100mg/kg BW). At 24-h post-implantation, fish were tethered to the seafloor and observed for behavioural metrics associated with predation. Blood samples were collected from a subset of fish to assess the physiological consequences of the implants. Cortisol- and RU486-implanted fish both had significantly higher plasma cortisol concentrations than sham fish, with blood glucose and plasma urea being elevated only in the former. Further, anti-predator behaviours and predation mortality did not differ significantly among treatments. Despite changes in physiological state, predation susceptibility was unaffected, a finding that may reflect the complex relationships linking the physiology and behaviour of an organism as well as potential tethering artefacts.

The energetic, physiological, and behavioral response of lemon sharks (Negaprion brevirostris) to simulated longline capture.

Commercial fisheries bycatch is a considerable threat to elasmobranch population recovery, and techniques to mitigate sub-lethal consequences can be improved with data on the energetic, physiological, and behavioral response of individuals to capture. This study sought to estimate the effects of simulated longline capture on the behavior, energy use, and physiological stress of juvenile lemon sharks (Negaprion brevirostris). Captive sharks equipped with acceleration biologgers were subjected to 1h of simulated longline capture. Swimming behaviors were identified from acceleration data using a machine-learning algorithm, energetic costs were estimated using accelerometer-calibrated relationships and respirometry, and physiological stress was quantified with point-of-care blood analyzers. During capture, sharks exhibited nine-fold increases in the frequency of burst swimming, 98% reductions in resting, and swam as often as unrestrained sharks. Aerobic metabolic rates during capture were 8% higher than for unrestrained sharks, and accounted for a 57.7% increase in activity costs when excess post-exercise oxygen consumption was included. Lastly, sharks exhibited significant increases in blood lactate and glucose, but no change in blood pH after 1h of capture. Therefore, these results provide preliminary insight into the behavioral and energetic responses of sharks to capture, and have implications for mitigating sub-lethal consequences of capture for sharks as commercial longline bycatch.

Hooking injury, physiological status and short-term mortality of juvenile lemon sharks (Negaprion bevirostris) following catch-and-release recreational angling.

Sport fishing for sharks, including fishing with the intent to release, is becoming more prevalent within the recreational angling community. Common targets of recreational anglers are juvenile lemon sharks (Negaprion brevirostris) that frequent shallow tropical nearshore habitats. In this study, we captured 32 juvenile lemon sharks (530-875 mm total length) with conventional angling gear (i.e. spinning rods, dead fish bait and 5/0 barbed circle hooks) from the coastal waters of Eleuthera, The Bahamas, to determine the consequences of capture for individual sharks. Each shark was examined for hooking injuries, blood sampled to quantify physiological disturbance, assessed for reflex impairment and then monitored to assess post-release behaviour and mortality. Four sharks (12.5%) died following release during the 15 min tracking period. Principal components (PC) analysis revealed four axes describing 66.5% of the variance for blood physiology parameters, total length and water temperature. The PC1 and PC3 scores, characterized by positive factor loadings for indicators of exercise-induced stress and blood ion concentrations, respectively, were significantly related to fight time but were not associated with short-term mortality. Short-term mortality was significantly related to factor scores for PC4 that loaded heavily for water temperature and total length. Ten sharks (31%) exhibited impaired reflexes, with loss of bite reflex being most prevalent. Sharks that died had the following characteristics: (i) they had two or more impaired reflexes; (ii) they were hooked in the basihyal; (iii) they exhibited no movement after the initial bout of directional swimming; and (iv) they experienced high water temperatures (i.e. >31°C). Collectively, these results indicate that for juvenile lemon sharks inhabiting tropical flats, fight time can influence the degree of physiological disturbance, while water temperature contributes to the likelihood of survival following release.

Use of portable blood physiology point-of-care devices for basic and applied research on vertebrates: a review.

Non-human vertebrate blood is commonly collected and assayed for a variety of applications, including veterinary diagnostics and physiological research. Small, often non-lethal samples enable the assessment and monitoring of the physiological state and health of the individual. Traditionally, studies that rely on blood physiology have focused on captive animals or, in studies conducted in remote settings, have required the preservation and transport of samples for later analysis. In either situation, large, laboratory-bound equipment and traditional assays and analytical protocols are required. The use of point-of-care (POC) devices to measure various secondary blood physiological parameters, such as metabolites, blood gases and ions, has become increasingly popular recently, due to immediate results and their portability, which allows the freedom to study organisms in the wild. Here, we review the current uses of POC devices and their applicability to basic and applied studies on a variety of non-domesticated species. We located 79 individual studies that focused on non-domesticated vertebrates, including validation and application of POC tools. Studies focused on a wide spectrum of taxa, including mammals, birds and herptiles, although the majority of studies focused on fish, and typical variables measured included blood glucose, lactate and pH. We found that calibrations for species-specific blood physiology values are necessary, because ranges can vary within and among taxa and are sometimes outside the measurable range of the devices. In addition, although POC devices are portable and robust, most require durable cases, they are seldom waterproof/water-resistant, and factors such as humidity and temperature can affect the performance of the device. Overall, most studies concluded that POC devices are suitable alternatives to traditional laboratory devices and eliminate the need for transport of samples; however, there is a need for greater emphasis on rigorous calibration and validation of these units and appreciation of their limitations.

Linking sensory biology and fisheries bycatch reduction in elasmobranch fishes: a review with new directions for research.

Incidental capture, or bycatch, in fisheries represents a substantial threat to the sustainability of elasmobranch populations worldwide. Consequently, researchers are increasingly investigating elasmobranch bycatch reduction methods, including some focused on these species' sensory capabilities, particularly their electrosensory systems. To guide this research, we review current knowledge of elasmobranch sensory biology and feeding ecology with respect to fishing gear interactions and include examples of bycatch reduction methods used for elasmobranchs as well as other taxonomic groups. We discuss potential elasmobranch bycatch reduction strategies for various fishing gear types based on the morphological, physiological, and behavioural characteristics of species within this diverse group. In select examples, we indicate how an understanding of the physiology and sensory biology of vulnerable, bycatch-prone, non-target elasmobranch species can help in the identification of promising options for bycatch reduction. We encourage collaboration among researchers studying bycatch reduction across taxa to provide better understanding of the broad effects of bycatch reduction methods.

The physiological response of the Caribbean reef shark (Carcharhinus perezi) to longline capture.

Longline fishing is the most common elasmobranch capture method around the world, yet the physiological consequences of this technique are poorly understood. To quantify the sub-lethal effects of longline capture in the commonly exploited Caribbean reef shark (Carcharhinus perezi), 37 individuals were captured using standard, mid-water longlines. Hook timers provided hooking duration to the nearest minute. Once sharks were landed, blood samples were taken and used to measure a suite of physiological parameters. Control data were obtained by sampling an additional three unrestrained Caribbean reef sharks underwater at an established shark feeding site. The greatest level of physiological disruption occurred after 120-180min of hooking, whereas sharks exposed to minimal and maximal hook durations exhibited the least disturbed blood chemistry. Significant relationships were established between hooking duration and blood pH, pCO(2), lactate, glucose, plasma calcium and plasma potassium. Longline capture appears more benign than other methods assessed to date, causing a shift in the stress response from acute at the onset of capture to a sub-acute regime as the capture event progresses, apparently facilitating a degree of physiological recovery. Continued investigation into the physiological response of elasmobranchs to longline capture is vital for the effective management of such fisheries.

The physiological response to anthropogenic stressors in marine elasmobranch fishes: a review with a focus on the secondary response.

Elasmobranchs (sharks, rays, and skates) are currently facing substantial anthropogenic threats, which expose them to acute and chronic stressors that may exceed in severity and/or duration those typically imposed by natural events. To date, the number of directed studies on the response of elasmobranch fishes to acute and chronic stress are greatly exceeded by those related to teleosts. Of the limited number of studies conducted to date, most have centered on sharks; batoids are poorly represented. Like teleosts, sharks exhibit primary and secondary responses to stress that are manifested in their blood biochemistry. The former is characterized by immediate and profound increases in circulating catecholamines and corticosteroids, which are thought to mobilize energy reserves and maintain oxygen supply and osmotic balance. Mediated by these primary responses, the secondary effects of stress in elasmobranchs include hyperglycemia, acidemia resulting from metabolic and respiratory acidoses, and profound disturbances to ionic, osmotic, and fluid volume homeostasis. The nature and magnitude of these secondary effects are species-specific and may be tightly linked to metabolic scope and thermal physiology as well as the type and duration of the stressor. In fishes, acute and chronic stressors can incite a tertiary response, which involves physiological changes at the organismal level, thereby impacting growth rates, reproductive outputs or investments, and disease resistance. Virtually no studies to date have been conducted on the tertiary stress response in elasmobranchs. Given the diversity of elasmobranchs, additional studies that characterize the nature, magnitude, and consequences of physiological stress over a broad spectrum of stressors are essential for the development of conservation measures. Additional studies on the primary, secondary, and tertiary stress response in elasmobranchs are warranted, with particular emphasis on expanding the range of species and stressors examined. Future studies should move beyond simply studying the effects of known stressors and focus on the underlying physiological mechanisms. Such studies should include the coupling of stress indicators with quantifiable aspects of the stressor, which will allow researchers to test hypotheses on survivorship and, ultimately, derive models that effectively link physiology to mortality. Studies of this nature are essential for decision-making that will result in the effective management and conservation of these species.