Predicting demography from species traits: larval development time and its sensitivity to warming depend on egg size in corals

doi:10.21203/rs.3.rs-5298759/v1

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Predicting demography from species traits: larval development time and its sensitivity to warming depend on egg size in corals

https://doi.org/10.21203/rs.3.rs-5298759/v1

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In the absence of demographic data, readily measurable life history traits, like egg size, can be used to predict how vital rates vary across species, facilitating modeling and analysis of high-diversity assemblages. We assessed the larval survival and competency dynamics of four previously unstudied coral species at current and warmed temperatures, and combined it with data for three other species to assess how well egg size predicts the sensitivity to warming of mortality and the minimum time to competency, both determinants of larval dispersal. Minimum time to competency increased with egg size; moreover, warming-induced reductions in time to competency were greater for species with larger eggs. In contrast, mortality rate and its response to warming were both independent of egg size. These findings show how assemblage-level responses to environmental change can be inferred for diverse communities and indicate how warming-induced changes in larval biology may reshape reef coral metacommunities.

Acropora tenuis

Acropora digitifera

broadcast spawners

climate change

Ctenactis echinata

larval competency

Lobophyllia corymbosa

meta-analysis

trait-based ecology

Predicting how species and metacommunities will respond to environmental change is a key challenge for ecology (Orr et al. 2020; Thompson et al. 2021). For relatively species-poor communities this can be accomplished using models of metacommunity dynamics calibrated by direct measurements of growth, intra- and interspecific competition and/or facilitation, reproduction, recruitment and dispersal (e.g., Pacala et al. 1996). Such models not only help us answer ecological and evolutionary questions, such as about the relative importance of different coexistence mechanisms (Adler et al. 2010), they can also be used to inform management decisions, such as the design of networks of protected areas (Born et al. 2008; Edwards et al. 2010). For species-rich communities, however, such demographic data are rarely available, nor easily collectable, for the vast majority of species (Urban et al. 2016). One solution to this is to characterize how the demographic characteristics of species vary with more readily measurable species traits (Adler et al. 2014; Madin et al. 2016; Wieczynski et al. 2021), and to infer species-level demography based on species’ portfolios of traits.

Egg size is an easy-to-measure, highly conserved life history trait (Robertson and Collin 2015) which, in many groups, can serve as proxy for larval development time, and consequently dispersal potential. Larger eggs tend to have greater energy content (McEdward and Morgan 2001; Robertson and Collin 2015) and produce larger feeding larvae or newly metamorphosed recruits (Strathmann 1977; Sinervo 1990; Bernardo 1996; Fox and Czesak 2000; Garcia-Barros 2000; Marshall and Keough 2008; Allen and Marshall 2014). Egg size is inversely related to larval development time (Berrill 1935; Steele 1977; Emlet et al. 1987; Garcia-Barros 2000), both within (Marshall and Bolton 2007) and among species (Staver and Strathmann 2002) with the same feeding modes and at similar temperature. Increased metabolic demands for oxygen, coupled with a decrease in the capacity to acquire oxygen by diffusion, have been hypothesized to explain this pattern (Einum et al. 2002). Because planktonic period affects dispersal potential, egg size can also be a good proxy for larval dispersal potential in some groups, particularly those whose larvae are weak swimmers (see Shanks 2009 for influence of larval swimming on homing behaviors that can limit dispersal). Dispersal is a particularly important demographic process because it influences species distribution, persistence, gene flow and local adaptation.

For stony corals, the main structure-formers of coral reef ecosystems, dispersal is limited to the larval stage. How long larvae remain alive and when they become able to settle are important determinants of coral larval dispersal patterns. However, corals are a highly diverse group, and the calibration of competence dynamics for all coral species in the metacommunity would be time-consuming and costly. In broadcast spawning corals, egg size explains ca. 80% of the variation in time to motility (i.e. the time it takes from egg fertilization to the acquisition of cilia and start of movement; Figueiredo et al. 2013); however, the potential relationship between egg size and the minimum time to competency (when larvae have developed the necessary structures to settle and metamorphose when exposed to suitable settlement cues) has not yet been assessed, nor has the relationship between egg size and mortality rate. Such relationships could facilitate inferences about assemblage-level changes in dispersal potential in reef corals, particularly if egg size explains a substantial proportion of the interspecific variation in these parameters.

Larval dispersal patterns are expected to be affected by climate change. Warming accelerates embryogenesis and larval development of most marine ectothermic species (Hoegh-Guldberg and Pearse 1995; O’Connor et al. 2007), including multiple broadcast spawning corals (Nozawa and Harrison 2007; Heyward and Negri 2010), changes that can alter larval dispersal potential. Specifically, local retention increases (Figueiredo et al. 2014) and reef connectivity decreases (Figueiredo et al. 2022) when development time is shorter. However, the magnitude of interspecific variability in such responses, and the role of modulators such as egg size, have not been assessed. In this study, we experimentally assessed the larval survival and competency dynamics of four previously unstudied coral species at current and warmed temperatures. We then combined these new data with previously-collected data from three other coral species in meta-regressions. Specifically, we estimate the magnitude of interspecific variation in these demographic quantities, we assess the extent to which key parameters driving larval dispersal potential are predicted by egg size, and we determine whether egg size predicts the magnitude of warming-induced changes in those parameters. Our findings highlight how changes to disperal dynamics might shape the metacommunity structure and dynamics of reef coral assemblages in the coming decades, and illustrate how readily-measurable trait values can inform projected changes to high-diversity communities more broadly.

Larval survival and competency experiments

Six colonies of Ctenactis echinata, Acropora digitifera, A. tenuis, and Lobophyllia corymbosa (identified following Veron 2000) were collected nearshore at Sesoko Island (Okinawa, Japan) before spawning in June or July 2013. After spawning in the laboratory at Sesoko Marine Research Station of University of the Ryukyus, the gametes of all colonies of the same species were collected and mixed. Fifty eggs of each species were measured under a stereomicroscope with a calibrated micrometer. Embryos/larvae of each species were reared at three temperatures: ambient, + 2 and + 4°C (respectively 27, 29 and 31°C), except for A. digitifera whose embryos/larvae were only reared at ambient and + 4°C, due to an insufficient number of larvae for an intermediate temperature treatment. Larval survival was assessed daily following an initial stock of four replicates of 50 embryos per temperature. For A. digitifera and L. corymbosa, the larval competency at each temperature was assessed using four replicates of 100 embryos. In each replicate, embryos were stocked to a 200mL glass jar filled with filtered (0.2 µm) seawater with a 2 × 2 × 0.5 cm pre-conditioned ceramic tile to induce settlement and metamorphosis. Every day, the tile was removed from the jar, censused under a scope for metamorphosed corals, and replaced by another tile; censuses were performed until the day no larvae were observed settling in any temperature treatment. The number of larvae that remained alive but had not settled until the end of the experiment was also counted. For C. echinata and A. tenuis, embryos were reared in three or more 2L bowls per temperature. To assess competency at each temperature, every day until day 6, and then on days 9, 16, 20, and 30 post-fertilization, four replicates of twenty larvae were haphazardly collected from all bowls and placed in 200mL glass jars filled with filtered seawater at the same temperature and provided a 2 × 2 × 0.5 cm pre-conditioned ceramic tile to induce settlement and metamorphosis. A day later the larvae that had settled and metamorphosed at each temperature were recorded.

Larval dynamics modelling

The larval survival and competency dynamics from seven species [four from this study (above) and three previously published (Figueiredo et al. 2014; Figueiredo et al. 2022)] representing a wide range of egg diameter [C. echinata (256µm), Cyphastrea japonica (297µm), Favites stylifera (454µm), A. tenuis (517µm), A. digitifera (521µm), A. millepora (540µm) and L. corymbosa (593µm)], were modelled using standard likelihood formulations, and parameters were estimated using maximum likelihood (Figueiredo et al. 2014; Figueiredo et al. 2022, detailed in Supplement). For each species, we used model selection to assess whether there was support for temperature-dependent mortality and competency rates by comparing a model where all parameters were independent of temperature with models where different combinations of parameters were dependent on temperature, using Akaike’s Information Criterion (AIC). The time to 25% mortality (M25) was estimnated as the time when \(\:1-{e}^{{\left(-\mu\:t\right)}^{v}}\)= 0.25. The variances of the minimum time to competency (t_c, i.e., the time it takes an embryo of broadcast spawning coral species to develop, settle and metamorphose when exposed to suitable settlement cues) and M25 were calculated by randomly generating 1000 (multivariate normal) of survival and competency model parameters [including temperature-(in)dependent log(t_c) and survival parameters log(µ) and log(ν)] using the variance-covariance matrix (i.e., inverse of the hessian matrix, evaluated at the maximum likelihood), then using each parameter set to generate 1000 estimates of the time to 25% mortality and t_c, and from these calculate their respective variance (Table 1).

Egg diameter effect on coral larval development and its sensitivity to warming

To describe the relationship between egg diameter and the minimum time to competency (t_c) at current temperature conditions in corals (Table 1), a mixed-effects metaregression was conducted using the maximum likelihood estimate of t_c and its estimated sampling variance for the same seven species, with species identity as a random factor and egg diameter (µm) as a moderator. To determine the effect of egg diameter on the minimum time to competency (t_c) at current temperature, this metaregression was compared with one where the egg diameter was not included as moderator using AIC and the Likelihood Ratio Test (LRT); both meta-regressions were run using the maximum likelihood method (as opposed to restricted maximum likelihood) to allow for the comparison of models with different fixed-effects structures (Zuur et al. 2009). Once the best model was identified, this was refit using restricted maximum likelihood to obtain less biased parameter estimates. The same methodology was used to test the effect of egg diameter on the time to 25% mortality (M25) at current temperature conditions.

To assess the efect of egg diameter on the warming-induced reduction of the minimum time to competency we used a two-step metaregression analysis. First, for each species, a fixed-effects metaregression of the minimum time to competency was conducted using the estimated minimum time to competency at each temperature and its variance (Table 1), and temperature as moderator. Then, a mixed-effects metaregression of the estimated slopes of each species’ metagression (Δt_c/ΔT, in h/°C) and their variance (calculated by squaring the product of the estimated slope by number of temperatures tested minus 1) was conducted using species identity as a random effect and egg diameter as the moderator. To assess the effect of egg diameter on the warming-induced changes to the minimum time to competency, this metaregression was compared, using the AIC and loglikelihood ratio test, with one where egg diameter was not included as moderator. The best model was refit using restricted maximum likelihood (REML). The total interspecific variance explained by the moderator was estimated as the proportional reduction in the variance component of the model with the moderator, relative to the model without it (using ML). The same methodology was used to test the effect of egg diameter on the warming-induced changes to the time to 25% mortality (M25).

Larval survival and competency dynamics

For the species examined experimentally in this study, warming accelerated the acquisition of competency (i.e. reduced minimum time to competency by third to half) in Acropora tenuis, A. digitifera, and Lobophyllia corymbosa, and increased larval mortality (ca. 25–33%) of Ctenactis echinata, A. digitifera and A. tenuis, particularly during the early stages of larval development (embryogenesis). In contrast, the larval competency dynamics of C. echinata and the larval survival of L. corymbosa were not affected by warming (Fig. 1, and Tables S1-4 and Figures S1-4 in Supplement).

Effect of egg diameter on coral larval development and its sensitivity to warming

Combining our estimates from the data above with those obtained from previous work in a meta-analysis revealed that the minimum time to competency (t_c) at current temperature (Table 1) significantly increased with egg diameter (taking ca. 19 h longer to develop per each 100 µm increase in egg diameter, LRT: p = 7×10^− 4, Fig. 2), with egg diameter explaining 80.5% of the total interspecific variance. Minimum time to competency decreased by about − 5.6 ± 1.5 h (mean ± SE) per each increased degree in temperature (°C); Δt_c/ΔT is normally distributed (One sample Kolmogorov-Smirnov test, D = 0.24, p = 0.75) and it varies considerably among species (SD = 3.6). The magnitude of Δt_c/ΔT significantly increased with egg diameter (loglikelihood ratio test, p = 0.02, Fig. 3), i.e. as temperature increased, species with larger egg diameter experienced a greater reduction in the minimum time to competency; 51.2% of the interspecific variance in Δt_c/ΔT was explained by egg diameter.

At current temperature, larvae took on average 96.3 ± 41.6 h (± SE) to reach 25% mortality (M25), and this time decreased by about − 2.4 ± 1.5 h on average (± SE) for every degree increase in temperature (°C) (Table 1). Both M25 and its sensitivity to warming (ΔM25/ΔT) varied substantially among species (SD = 3.6 and SD = 101.9, respectively). Model selection did not favor including egg diameter as a predictor of the time until 25% mortality at current temperature (loglikelihood ratio test, p = 0.36), nor did egg size predict the sensitivity of increase mortality to temperature (LRT, p = 0.22), suggesting it explains, at most, a small proportion of the total interspecific heterogeneity in mortality.

Table 1

Mean egg diameter (µm) and minimum time to competency (h) and time to 25% mortality and its estimated variances for each temperature for the seven species used in the metaregressions. * indicates species which larval dynamics were previously published in Figueiredo et al. (2014).
Species	Egg diameter (µm)	Minimum time to competency (h) (variance)			Time to 25% mortality (h) (variance)
Species	Egg diameter (µm)	27°C (ambient)	29°C (+ 2°C)	31°C (+ 4°C)	27°C (ambient)	29°C (+ 2°C)	31°C (+ 4°C)
Ctenactis echinata	256	28.35 (13.06)	28.35 (13.06)	28.35 (13.06)	4 (1.46)	2 (0.30)	1 (0.02)
Cyphastrea japonica*	297	50.00 (2.11)	42.63 (4.44)	41.86 (5.49)	44 (42.02)	39 (37.18)	12 (12.01)
Favites stylifera*	454	60.04 (0.20)	51.57 (1.70)	44.12 (3.41)	194 (538.5)	194 (538.5)	194 (538.5)
Acropora tenuis	517	85.32 (4.30)	60.27 (1.09)	41.66 (0.11)	1659 (282476)	647 (22778)	361 (5527)
Acropora digitifera	521	97.00 (1.71)		81.88 (0.09)	305 (7351.63)		30 (89.95)
Acropora millepora*	540	103.81 (1.08)	86.34 (0.25)	62.67 (0.19)	18 (13.95)	5 (1.62)	3 (0.42)
Lobophyllia corymbosa	593	83.26 (1.30)	79.02 (3.07)	55.14 (2.90)	57 (41.46)	57 (41.46)	57 (41.46)

In this study, we identified strong relationships between egg size and minimum time to competency in a number of broadcast spawning corals. These relationships suggest that a substantial proportion of the interspecific variation in larval competency dynamics, and how this is affected by warming, is predictable from readily measurable characteristics of the eggs. Specifically, the minimum time to competency increases with egg diameter, making the larvae of species with larger eggs slower to develop. Additionally, species with larger eggs also experienced more substantial reductions of the minimum time to competency with increasing temperatures. As a result, broadcasting spawning coral species with larger eggs are expected to endure greater changes to their dispersal patterns than species with smaller eggs. The estimated trait-based relationships provided by this study, including the distribution of the warming-induced increase in mortality, the relationship between egg diameter and minimum time to competency, and the relationship between egg diameter and warming-induced changes to the minimum time to competency, can therefore facilitate projections of metacommunity scale responses to environmental change.

The diameter of the eggs of broadcast spawning corals is a good predictor of their minimum time to competency at current temperature conditions (Fig. 2) likely because larger eggs have lower respiratory exchange and thus longer larval development times. From marine to aquatic environments, within closely related animals which inhabit the same temperature regime, species with larger eggs typically take longer to complete larval development (Staver and Strathmann 2002). Egg size has been found to be a good proxy for larval development time in multiple groups, including amphipods (Steele and Steele 1973), ascidians (Berril 1935; Staver and Strathmann 2002), bivalves (Staver and Strathmann 2002), butterflies (Garcia-Barros 2000), copepods (McLaren 1966), decapods (Wear 1974), fish (Duarte and Alcaraz 1989), frogs (McLaren and Cooley 1972), gastropods (Staver and Strathmann 2002), hydromedusas (Staver and Strathmann 2002), nudibranchs (Wray and Hadfield 1987) and seastars (Strathmann and Staver 2002). This relationship between egg size and larval development time is explained by the ratio of the surface-area to the volume of the egg, as this determines the rate of respiratory exchange and thus control development rate (Berril 1935). However, it is not oxygen diffusion that limits larval development as during early stages development does not require much oxygen, and oxygen consumption increases relatively slowly with increasing egg mass (Berril 1935; Einum et al. 2002). Interestingly, it appear to be driven by carbon dioxide, which can be very toxic and thus needs to be removed from the egg. As egg size (and ratio of the surface-area to the volume) increases, the elimination of CO₂ from the egg and embryo surfaces is progressively harder, requiring a reduction in the pace of larval development so that CO₂ generation does not outpace its elimination (Berril 1935).

The minimum time to competency of the larvae of broadcast spawning coral species decreases as temperatures warm (Table 1, Fig. 1) because the latter accelerates metabolic rates and consequently larval development. Temperature is of greatest influence upon the duration of larval development (Thorson 1950). Within the species’ tolerance range, low temperatures lengthen the pelagic life, while high temperatures shorten it (provided food availability or maternal reserves are high enough to support the increased metabolic rate at these high temperatures) (Berril 1935; O’Connor et al. 2007). This trend has been reported in multiple marine and freshwater animals, such as amphipods (Steele and Steele 1973), ascidians (Berril 1935), brine shrimp (Figueiredo et al. 2009), copepods (McLaren 1966), decapods (Wear 1974; Kumlu et al. 2000, Figueiredo and Narciso 2006), fish (Duarte and Alcaraz 1989), frogs (McLaren and Cooley 1972), sea cucumber (Asha and Muthiah 2005), and other corals (Nozawa and Harrison 2007; Randall and Szmant 2009a,b; Heyward and Negri 2010), and is commonly exploited by aquaculture facilities to accelerate production (Rothlisberg 1998; Morais et al. 2014). Warmer temperature increases the activity of enzymes, accelerating fundamental biochemical processes and consequently metabolic rates (Clarke and Fraser 2004) and larval development. Together with previous observations, the additional warm-induced larval dynamics provided in this study confirm that the hastening of larval development under warmer conditions likely extends to all broadcasting spawning coral species. The results of this study suggest that estimates of the change in minimum time to competency with increased temperature can be randomly generated from a normal distribution (mean=-5.6 and SD = 3.6). The ability to produce estimates of the minimum time coral larvae remain in the water column before being able to settle without information on the reproductive ecology of the species in a community can improve estimates of demographic connectivity of coral populations. However, these estimates can be even further improved if the egg size of the coral species present in a community are known.

The extent of the reduction of minimum time to competency under warmer temperatures is directly related to egg size, with species with larger eggs experiencing greater reductions. Coral species with larger eggs experience a greater reduction in (absolute) time to competency likely as a result from having a longer larval development, and thus any acceleration in development shortens considerably more the time required for their larvae to become competent. The results of this study suggest that the decrease in the minimum time to competency of coral larvae, Δt_c (h), with temperature increase, ΔT (°C), increases with the mean egg diameter of the species (µm), and can be described by the following (Fig. 3):

Δt_c/ΔT (h/°C) = 5.385–2.38×10^− 2 × Egg Diameter (µm)

For each degree (°C) temperature increases, species with larger eggs (> 450µm) will become ready to settle 5-10h earlier, while species with smaller eggs (< 300µm) will only develop 1-2h faster. The greater the increase in temperature, the more striking these differences become. For example, a 4°C increase leads to a hastening of development of 0-8h for species with small eggs vs. 20-40h for species with large eggs. This magnitude of shortening of larval development (1 day or more) can cause a substantial change in coral larval dispersal patterns and connectivity, namely an increase in local retention, self-recruitment and weakening of connectivity (Figueiredo et al. 2022), with important consequences for the persistence and recovery of populations following disturbances (Figueiredo et al. 2022), as well as genetic diversity and adaptation processes. Since the experiments used to estimate this relationship did not expose the adult coral colonies to warmer conditions, the proposed equation presumes that their egg diameter will not be altered by warming. Exposure to warming conditions has been found to decrease the egg size in several ectotherms (Atkinson et al. 2001; Robertson and Collin 2015; Ly and Collin 2021), however there are many exceptions to this trend (Chambers 1997; Fox and Czesak 2000: Atkinson et al. 2001). The effect of temperature on the egg size of broadcasting spawning corals remains poorly studied. Existing studies suggest that egg size may vary with some environmental conditions, but temperature stress seems to have a minor effect on egg size. Corals associated with the more thermally tolerant algal Durisdinium symbionts produced eggs with a lower lipid content and smaller size than the ones associated with the thermally-sensitive Cladocopium; however, while exposure of adult corals to warming also reduced the number and lipid content of the eggs produced, it did not affect the size of their eggs (Jones and Berkelmans 2011). Temperature differences between seasons have also been found not alter the egg size nor fecundity of four biannually spawning coral species (Foster and Gilmour 2020). This evidence is still insufficient to determine if and to which extent the egg size of coral species will be altered by warming. However, if the egg size of corals does shrink under warmer temperatures, since egg size is directly related to the minimum time to competency, the reduction of time to competency under warm conditions described above would be even more pronounced than predicted here.

For most species studied, larval mortality increased with temperature, regardless of egg size. Increased mortality during embryogenesis and larval development at higher (than normal/optimal) temperatures has been reported, to a greater or lower extent, in most ectotherms (e.g. echinoderms [Byrne et al. 2013], fish [Madeira et al. 2016], and crustaceans [Quinn 2017]). This trend is often explained by an acceleration of enzyme activity under warmer temperatures that accelerates cell division, and in the process potentially increases the chances of malformation due to lack of synchronicity among tissues. While the larvae of most coral species suffer higher mortality under higher temperatures (Nozawa and Harrison 2007; Randall and Szmant 2009a,b; Heyward and Negri 2010; Figueiredo et al. 2014; Putnam andGates 2015; Graham et al. 2017; Matsuda et al. 2021; Figueiredo et al. 2022, this study), we did not find a relationship between mortality and egg size. It is likely that even if egg size partially explains the differential mortality among coral species, its effect will be much smaller than the effects of parental genotype, history of stress exposure, food quantity and quality, and/or symbiont community (Jones and Berkelmans 2011; Baums et al. 2013; Padilla-Gamiño et al. 2013; Kirk et al. 2018).

Coral species with larger eggs are expected to disperse further but, as oceans warm, experience greater reductions in dispersal distance and connectivity. Since coral larvae are mostly passive dispersers (Hata et al. 2017), the direct relationship between egg size and time to competency, and of the latter with dispersal potential (Figueiredo et al. 2013, 2014, 2022), means species with larger eggs have a greater potential to disperse away from their parental habitat. Furthermore, as the reduction of the minimum time to competency with warming is greater in species with larger eggs, we project that as oceans warm, these species will experience greater reductions in dispersal distance and connectivity. Reducing the time larvae spend in the plankton (pre-competency time) will also increase the probability larvae will settle before being flushed from their natal reef, thus experiencing a greater relative increase in local retention and self-recruitment. The broadcast spawning corals with the largest eggs are from the genera Lobophyllia and Acropora. Acropora is the most diverse and abundant genus of reef-building corals (Veron 2000); their branching morphology provides habitat to a multitude of reef fish species. An increase in local retention and self-recruitment, and reduction in connectivity of these species is expected to reduce the inter-reef genetic exchange, increasing isolation and reducing gene flow and recruitment subsidies following disturbance. Considering Acropora are also some of the most susceptible species to bleaching (Marshall and Baird 2000), these changes are likely to considerably reshape meta-communities in the Indo-Pacific.

Measuring demographic parameters to predict assemblage-level responses to environmental change in highly diverse communities is extremely challenging, but readily measurable life history traits, like egg size, may at least in some cases be used to characterize interspecific differences, and thus facilitate modeling and analysis and predictions. Existent projections of how demographic parameters, and consequently species abundance and distribution, will be affected by environmental change use empirically-calibrated models or the ecological niche of the species based on time-series distributions over large spatial scales (e.g. O’Connor et al. 2007 [marine fish and invertebrates]; Buckley et al. 2011 [butterflies]; Bowler et al. 2017 [many groups]; Sanchez-Salguero et al. 2017 [trees]; Simon-Nutbrown et al. 2020 [coralline algae]; Benedetti et al. 2021 [plankton]). For corals, projections of warm-induced changes to larval dispersal patterns and connectivity have been based on “generic” larval dynamics (Wood et al. 2016), thus not reflecting the inter-specific variability, or focused on a single species (Figueiredo et al. 2022; Holstein et al. 2022); projections for entire coral communities are inexistent. Experimentally assessing how demographic parameters of a species will change with warming often requires access to a temperature-controlled laboratory, is time-consuming, labor-intensive and expensive; for larval development dynamics, data collection is often restricted to one annual reproductive event. Doing experimental calibrations for all species of a diverse taxonomic group, such as corals, is virtually impossible. The relationships between readily-measurable life history traits and effects of environmental change on demographic parameters, such as the one described here for corals, allow us to bypass that, facilitating estimations of future abundance and ability to recover from disturbances for unstudied species and, more importantly, entire meta-communities. For corals, projections of future dispersal using the relationship between egg size and warm-driven changes in larval development time would greatly benefit the design of networks of marine protected areas and siting of reef restoration efforts that aim to counteract the negative effects of climate change. The relationship between egg size and warming-effects on larval development has not been studied in other taxonomic groups, thus it is not clear if the species with larger eggs within other taxonomic groups are also expected to experience greater declines in dispersal. However, considering (within taxonomic group and habitat) relationships between egg size and larval development time (Berrill 1935 [tunicates]; McLaren 1966 [copepods]; McLaren and Cooley 1972 [frogs]; Steele and Steele 1973 [amphipods]; Steele and Streele 1975 [crustaceans]; Wray and Hadfield 1987 [opisthobranchs (Nudibranchia)]; Duarte and Alcaraz 1989 [marine and freswater fish]; Garcia-Barros 2000 [butterflies]), and the known relationship between temperature and larval developmental rates (Hoegh-Guldberg and Pearse 1995 [asteroids]; O’Connor et al. 2007 [marine fish and invertebrates]), it is quite possible that that is the case. Further studies have the potential greatly expand our ability to predict the dispersal, and consequently the persistence, of large multi-taxon communities.

Funding:

This study was supported by the Australian Research Council (DP110101168 to A.H.B. and J.F and DP0880544 to S.R.C), Japan Society for the Promotion of Science (Fellowship to J.F., hosted by S.H.) and the State of Queensland (Smart Futures Fellowship to J.F.).

Conflict of interest:

J.F. serves as a Topic Editor for Coral Reefs. Authors declare no other conflict of interests.

Author Contribution

J.F., A.B. and S.C. designed the study. J.F., A.B. and S.H. collected data. J.F. and S.C. performed modeling work, the meta-analysis and analyzed results. J.F. wrote the first draft of the manuscript. All authors contributed substantially to revisions.

Acknowledgements:

The Smithsonian Tropical Research Institute in Panama hosted a sabbatical visit from J.F. to conduct the analysis for this study.

Data Availability

Data is provided within the manuscript or supplementary information and/or from the authors.

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Competing interest reported. J.F. serves as a Topic Editor for Coral Reefs.

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Predicting demography from species traits: larval development time and its sensitivity to warming depend on egg size in corals

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Introduction

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Larval survival and competency experiments

Larval dynamics modelling

Egg diameter effect on coral larval development and its sensitivity to warming

Results

Larval survival and competency dynamics

Effect of egg diameter on coral larval development and its sensitivity to warming

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