Body mass, temperature, and depth shape the maximum intrinsic rate of population increase in sharks and rays.

An important challenge in ecology is to understand variation in species' maximum intrinsic rate of population increase, r max , not least because r max underpins our understanding of the limits of fishing, recovery potential, and ultimately extinction risk. Across many vertebrate species, terrestrial and aquatic, body mass and environmental temperature are important correlates of r max . In sharks and rays, specifically, r max is known to be lower in larger species, but also in deep sea ones. We use an information-theoretic approach that accounts for phylogenetic relatedness to evaluate the relative importance of body mass, temperature, and depth on r max . We show that both temperature and depth have separate effects on shark and ray r max estimates, such that species living in deeper waters have lower r max . Furthermore, temperature also correlates with changes in the mass scaling coefficient, suggesting that as body size increases, decreases in r max are much steeper for species in warmer waters. These findings suggest that there are (as-yet understood) depth-related processes that limit the maximum rate at which populations can grow in deep-sea sharks and rays. While the deep ocean is associated with colder temperatures, other factors that are independent of temperature, such as food availability and physiological constraints, may influence the low r max observed in deep-sea sharks and rays. Our study lays the foundation for predicting the intrinsic limit of fishing, recovery potential, and extinction risk species based on easily accessible environmental information such as temperature and depth, particularly for data-poor species.

Sharkipedia: a curated open access database of shark and ray life history traits and abundance time-series.

A curated database of shark and ray biological data is increasingly necessary both to support fisheries management and conservation efforts, and to test the generality of hypotheses of vertebrate macroecology and macroevolution. Sharks and rays are one of the most charismatic, evolutionary distinct, and threatened lineages of vertebrates, comprising around 1,250 species. To accelerate shark and ray conservation and science, we developed Sharkipedia as a curated open-source database and research initiative to make all published biological traits and population trends accessible to everyone. Sharkipedia hosts information on 58 life history traits from 274 sources, for 170 species, from 39 families, and 12 orders related to length (n = 9 traits), age (8), growth (12), reproduction (19), demography (5), and allometric relationships (5), as well as 871 population time-series from 202 species. Sharkipedia relies on the backbone taxonomy of the IUCN Red List and the bibliography of Shark-References. Sharkipedia has profound potential to support the rapidly growing data demands of fisheries management, international trade regulation as well as anchoring vertebrate macroecology and macroevolution.

Estimating marine survival of Atlantic salmon using an inverse matrix approach.

The marine phase of anadromous Atlantic salmon (Salmo salar) is the least known yet one of the most crucial with regards to population persistence. Recently, declines in many salmon populations in eastern Canada have been attributed to changes in the conditions at sea, thus reducing their survival. However, marine survival estimates are difficult to obtain given that many individuals spend multiple winters in the ocean before returning to freshwater to spawn; therefore, multiple parameters need to be estimated. We develop a model that uses an age-structured projection matrix which, coupled with yearly smolt and return abundance estimates, allows us to resample a distribution of matrices weighted by how close the resulting return estimates match the simulated returns, using a sample-importance-resampling algorithm. We test this model by simulating a simple time series of salmon abundances, and generate six different scenarios of varying salmon life histories where we simulate data for one-sea-winter (1SW)-dominated and non-1SW dominated populations, as well as scenarios where the proportion returning as 1SW is stable or highly variable. We find that our model provides reasonable estimates of marine survival for the first year at sea (S1), but highly uncertain estimates of proportion returning as 1SW (Pr) and survival in the second year at sea (S2). Our exploration of variable scenarios suggests the model is able to detect temporal trends in S1 for populations that have a considerable 1SW component in the returns; the ability of the model to detect trends in S1 diminishes as the proportion of two-sea-winter fish increases. Variability in the annual proportion of fish returning as 1SW does not seem to impact model accuracy. Our approach provides an instructive stepping-stone towards a model that can be applied to empirical abundance estimates of Atlantic salmon, and anadromous fishes in general, and therefore improve our knowledge of the marine phase of their life cycles as well as examining spatial and temporal trends in their variability.

Population productivity of shovelnose rays: Inferring the potential for recovery.

There is recent evidence of widespread declines of shovelnose ray populations (Order Rhinopristiformes) in heavily fished regions. These declines, which are likely driven by high demand for their fins in Asian markets, raises concern about their risk of over-exploitation and extinction. Using life-history theory and incorporating uncertainty into a modified Euler-Lotka model, the maximum intrinsic rates of population increase (rmax) were estimated for nine species from four families of Rhinopristiformes, using four different natural mortality estimators. Estimates of mean rmax, across the different natural mortality methods, varied from 0.03 to 0.59 year-1 among the nine species, but generally increased with increasing maximum size. Comparing these estimates to rmax values for other species of chondrichthyans, the species Rhynchobatus australiae, Glaucostegus typus, and Glaucostegus cemiculus were relatively productive, while most species from Rhinobatidae and Trygonorrhinidae had relatively low rmax values. If the demand for their high-value products can be addressed then population recovery for some species is likely possible, but will vary depending on the species.

Ecological lifestyles and the scaling of shark gill surface area.

Fish gill surface area varies across species and with respect to ecological lifestyles. The majority of previous studies only qualitatively describe gill surface area in relation to ecology and focus primarily on teleosts. Here, we quantitatively examined the relationship of gill surface area with respect to specific ecological lifestyle traits in elasmobranchs, which offer an independent evaluation of observed patterns in teleosts. As gill surface area increases ontogenetically with body mass, examination of how gill surface area varies with ecological lifestyle traits must be assessed in the context of its allometry (scaling). Thus, we examined how the relationship of gill surface area and body mass across 11 shark species from the literature and one species for which we made measurements, the Gray Smoothhound Mustelus californicus, varied with three ecological lifestyle traits: activity level, habitat, and maximum body size. Relative gill surface area (gill surface area at a specified body mass; here we used 5,000g, termed the 'standardized intercept') ranged from 4,724.98 to 35,694.39 cm2 (mean and standard error: 17,796.65 ± 2,948.61 cm2 ) and varied across species and the ecological lifestyle traits examined. Specifically, larger-bodied, active, oceanic species had greater relative gill surface area than smaller-bodied, less active, coastal species. In contrast, the rate at which gill surface area scaled with body mass (slope) was generally consistent across species (0.85 ± 0.02) and did not differ statistically with activity level, habitat, or maximum body size. Our results suggest that ecology may influence relative gill surface area, rather than the rate at which gill surface area scales with body mass. Future comparisons of gill surface area and ecological lifestyle traits using the quantitative techniques applied in this study can provide further insight into patterns dictating the relationship between gill surface area, metabolism, and ecological lifestyle traits.

An Introduction to Modelling Abundance and Life History Parameters in Shark Populations.

Elasmobranchs play critically important ecological roles throughout the world's oceans, yet in many cases, their slow life histories and interactions with fisheries makes them particularly susceptible to exploitation. Management for these species requires robust scientific input, and mathematical models are the backbone of science-based management. In this chapter, we provide an introductory overview of the use of mathematical models to estimate shark abundance. First, we discuss life history models that are used to understand the basic biology of elasmobranchs. Second, we cover population dynamics models, which are used to make inferences regarding population trend, size, and risk of extinction. Finally, we provide examples of applied models used to assess the status of elasmobranchs in the Northeast Pacific Ocean to guide management for these species. This chapter is not a comprehensive review of quantitative methods, but rather introduces various mathematical tools in fisheries management, with a focus on shark management in the Northeast Pacific Ocean.

Growth, productivity, and relative extinction risk of a data-sparse devil ray.

Devil rays (Mobula spp.) face intensifying fishing pressure to meet the ongoing international demand for gill plates. The paucity of information on growth, mortality, and fishing effort for devil rays make quantifying population growth rates and extinction risk challenging. Furthermore, unlike manta rays (Manta spp.), devil rays have not been listed on CITES. Here, we use a published size-at-age dataset for the Spinetail Devil Ray (Mobula japanica), to estimate somatic growth rates, age at maturity, maximum age, and natural and fishing mortality. We then estimate a plausible distribution of the maximum intrinsic population growth rate (rmax) and compare it to 95 other chondrichthyans. We find evidence that larger devil ray species have low somatic growth rate, low annual reproductive output, and low maximum population growth rates, suggesting they have low productivity. Fishing rates of a small-scale artisanal Mexican fishery were comparable to our estimate of rmax, and therefore probably unsustainable. Devil ray rmax is very similar to that of manta rays, indicating devil rays can potentially be driven to local extinction at low levels of fishing mortality and that a similar degree of protection for both groups is warranted.

Diagnosing the dangerous demography of manta rays using life history theory.

Background. The directed harvest and global trade in the gill plates of mantas, and devil rays, has led to increased fishing pressure and steep population declines in some locations. The slow life history, particularly of the manta rays, is cited as a key reason why such species have little capacity to withstand directed fisheries. Here, we place their life history and demography within the context of other sharks and rays. Methods. Despite the limited availability of data, we use life history theory and comparative analysis to estimate the intrinsic risk of extinction (as indexed by the maximum intrinsic rate of population increase r max) for a typical generic manta ray using a variant of the classic Euler-Lotka demographic model. This model requires only three traits to calculate the maximum intrinsic population growth rate r max: von Bertalanffy growth rate, annual pup production and age at maturity. To account for the uncertainty in life history parameters, we created plausible parameter ranges and propagate these uncertainties through the model to calculate a distribution of the plausible range of r max values. Results. The maximum population growth rate r max of manta ray is most sensitive to the length of the reproductive cycle, and the median r max of 0.116 year(-1) 95th percentile [0.089-0.139] is one of the lowest known of the 106 sharks and rays for which we have comparable demographic information. Discussion. In common with other unprotected, unmanaged, high-value large-bodied sharks and rays the combination of very low population growth rates of manta rays, combined with the high value of their gill rakers and the international nature of trade, is highly likely to lead to rapid depletion and potential local extinction unless a rapid conservation management response occurs worldwide. Furthermore, we show that it is possible to derive important insights into the demography extinction risk of data-poor species using well-established life history theory.