Population dynamics of two small pelagic fish in the central-south area off Chile: delayed density-dependence and biological interaction

It is widely believed that environmental variability is the main cause for fluctuations in commercially exploited small pelagic fish populations around the world. Nevertheless, density-dependent factors also can drive population dynamics. In this paper, we analyzed thirteen years of a relative abundance index of two clupeoids fish populations coexisting in the central-south area off Chile, namely the common sardine, Strangomera bentincki, and anchovy, Engraulis ringens. We applied the classical diagnostic tools of time series analysis to the observed time-series. Also, the realized per capita population growth rate was studied with the aim of detecting the feedback structure that is characterizing the population dynamics of the two species. The analysis suggests that population fluctuations of the two species have an important density-dependent component, displaying first-order (direct density-dependent) and second-order (delayed density-dependent) simultaneously. The density-dependent component explained 70.5 and 55.6 % of the realized per capita population growth rate of common sardine and anchovy, respectively. The deterministic skeleton model showed an asymptotic convergence to equilibrium density. In presence of a stochastic environment, fluctuations were reproduced for the species showing a component of fluctuation with a period of 4 year. The intrinsic dynamics of each species is typical of interacting species resulting from trophic interactions. It is postulated that the second-order dynamics of S. bentincki and E. ringens in central-south Chile, may be the result from interactions with a specialist predator (the fishing fleet), interacting with exogenous environmental factors.     

The complexation chemistry of cyclohexaamyloses: Adducts with 1-adamantanecarboxylic acid and anion

A study is reported of complexation reactions of cyclohexaamylose (Cy) with 1-adamantanecarboxylic acid and its anion using conductometry, pH potentiometry, and ¹³C nmr spectrometry. Binary and ternary (2 mol Cy/mol substrate) complexes are detected with both the acid and anion, and standard entropies and enthalpies of complexation are determined from the temperature dependences of the formation constants for all except the very weak ternary complex with the anion. Both the ¹³C nmr results and the entropy of complexation confirm the earlier suggestion that the anion in binary complexation is structured with the adamantanyl group in proximity to, but not penetrating, the Cy cavity. However, a negative ΔS^o for formation of this complex is reported which casts doubt on an earlier proposal that the adamantyl binary complex binding mode involves an “apolar” mechanism accompanied by loss of solvated water molecules. Values are also reported for pK_a, ΔH^o, and ΔS^o for the aqueous dissociation of 1-adamantanecarboxylic acid.     

The relationship between nutrient feed rate and specific growth rate in fed batch cultures

Summary Equations are described which relate nutrient feed rate to specific microbial growth rate in fed batch culture. Fed batch cultures are classified into three types: 1) those allowing constant specific microbial growth rate, 2) those in which the rate of change of flow rate is constant and 3) those in which the nutrient flow rate is constant. The basic properties of these three types are described.Symbols F medium flow rate, L³ T^–1 - F _o medium flow rate at zero time, L³ T^–1 - g rate of change of flow rate with time, L³ T^–2 - K _v volume constant, being the total cell weight at zero time divided by the product of the yield coefficient and growth-limiting substrate concentration in the feed, L³ - s _r growth limiting substrate concentration in the feed, ML^–3 - V volume of liquid in the growth vessel, L³ - V _f volume of medium fed to the growth vessel, L³ - V _o volume of liquid in the growth vessel at zero time, L³ - X total weight of cells, M - x concentration of cells, ML^–3 - X _g total weight of cells grown, M - X _o total weight of cells at zero time, M - Y yield coefficient, weight of cells grown per unit weight of growth-limiting substrate - specific microbial growth rate, T^–1     

On the interpretation of some planktonology equations

Summary The concept of the mean value of a function is used to interpret some population-dynamics equations. The well-known formula for the per capita growth rate r gives a precise mean value for any (not only exponentially growing) populations. This result is used to derive the birth and death rate equations of Paloheimo (1974) with minimal initial limitations.Abbreviations t time - N number of animals - E number of eggs - r specific (i.e. per capita) population growth rate - b specific birth rate - d specific death rate - D duration of embryonic development     

Perinatal adaptation in mammals: the impact of metabolic rate

Mammalian birth is accompanied by profound changes in metabolic rate that can be described in terms of body size relationship (Kleiber's rule). Whereas the fetus, probably as an adaptation to the low intrauterine pO₂, exhibits an “inappropriately” low, adult-like specific metabolic rate, the term neonate undergoes a rapid metabolic increase up to the level to be expected from body size. A similar, albeit slowed, “switching-on” of metabolic size allometry is found in human preterm neonates whereas animals that are normally born in a very immature state are able to retard or even suppress the postnatal metabolic increase in favor of weight gain and O₂ supply. Moreover, small immature mammalian neonates exhibit a temporary oxyconforming behavior which enhances their hypoxia tolerance, yet is lost to the extent by which the size-adjusted metabolic rate is “locked” by increasing mitochondrial density. Beyond the perinatal period, there are no other deviations from metabolic size allometry among mammals except in hibernation where the temporary “switching-off” of Kleiber's rule is accompanied by a deep reduction in tissue pO₂. This gives support to the hypothesis that the postnatal metabolic increase represents an “escape from oxygen” similar to the evolutionary roots of mitochondrial respiration, and that the overall increase in specific metabolic rate with decreasing size might contribute to prevent tissues from O₂ toxicity.     

Effects of internal and external sodium on the sodium current-voltage relationship in thesquid giant axon

Summary The early transient current-voltage relationship was measured in internally perfused voltage clamped squid giant axons with various concentrations of sodium on the two sides of the membrane. In the absence of sodium on either side there is an outward transient current which is blocked by tetrodotoxin and varies with internal potassium concentration. The current increases linearly with voltage for positive potentials. Adding sodium ions internally increases the slope of the current-voltage relationship. Adding sodium ions externally also increases the slope between +10 and +80 mV. Adding sodium to both sides produces the sum of the two effects.The current-voltage relationships were fit by straight lines between +10 and +80 mV. Plotting the extrapolated intercepts with the current axis against the differences in sodium concentrations gave a straight line,I _o =–P(c _o –c _i )F.P, the Fickian permeability, is about 10^–4 cm/sec. Plotting the slopes in three dimensions against the two sodium concentrations gave a planeg=g _o +(aNa _o +bNa _i )F.a is about 10^–6 cm/mV-sec andb about 3×10^–6 cm/mV-sec. Thus the current-voltage relationship for the sodium current is well described byI=–P(c _o –c _i )F+(ac _o +bc _i )FV for positive potentials. This is the linear sum of Fick's Law and Ohm's Law.P/(a+b)=25±1 mV (N=6) and did not vary with the absolute magnitude of the currents. Within experimental error this is equal tokT/e orRT/F.Increasing temperature increasedP, a andb proportionately. Adding external calcium, lithium, or Tris selectively decreasedP anda without changingb. In the absence of sodium, altering internal and external potassium while observing the early transient currents suggests this channel is more asymmetric in its response to potassium than to sodium.     

Aspects of life-table studies and functional response of Lysiphlebia mirzai

The fecundity, reproductive rate, and survival of Lysiphlebia mirzai parasitising third instar nymphs of the cereal aphid Rhopalosiphum maidis were measured at six different host densities under constant laboratory conditions. The survival rate (l_x) of the female parasitoids was unaffected by host density, with an average adult life-span of 5–6 days at all densities. The age-specific fecundity rate (m_x) was host density-dependent. The value of m_x decreased rapidly from the first day of parasitisation. The number of hosts available determined the maximum possible number of mummies. At 200 hosts available per day, the average fecundity was 184.6 mummies/female; the maximum number of mummies yielded by any female was 200. The relationship between host density and the number of aphids parasitised per female was linear at 50 aphids/cage/day, but at higher host densities (100 aphids/cage/day) a significant curvilinear regression was observed. The intrinsic rate of natural increase (r_m) increased with increasing host density. Maximum value of r_m (0.262) was obtained at a host density of 200. The response of r_m to changes in host density and parasitoid sex ratio is shown. A typical type II functional response was observed for L. mirzai. The curve was described by a logistic curve, N_p=200/[1+exp(5.65–1.60 ln N_o)]. The search rate of the parasitoid was inverse host density-dependent. No significant variation in the sex ratio of F₁ offspring was observed at different initial host densities. Sex ratio values exceeded 0.5 at all host densities. The results evaluated the reproductive potential of L. mirzai as a promising biological control agent.     

Lactose-H(OH) transport system of Escherichia coli Multistate gated pore model based on half-sites stoichiometry for high-affinity substrate binding in a symmetrical dimer

A model is proposed for the d-galactoside-H⁺(⁻OH) transporter of Escherichia coli that accounts for essentially all the experimental observations established for this system to date. In this model, the functional unit is postulated to be a dimer (consisting of two copies of lacY-specified polypeptide) which spans the membrane with a 2-fold symmetry axis in the membrane plane (Lancaster, J.R. (1978) J. Theor. Biol. 75, 35–50). The functional dimer is assumed to possess a single pore flanked by an inner gate (g_i) and an outer gate (g_o) and encompassing two oppositely oriented galactoside binding sites, designated m and μ. When g_o is open and g_i is closed under non-energized conditions, binding site m adopts a configuration defined as State A (i.e., m_o^A) exhibiting high affinity toward Class G^a galactosides (thiodigalactoside, melibiose, α-p-nitrophenylgalactoside) but low affinity for Class G^b galactosides (lactose, β-o-nitrophenylgalactoside, β-isopropylthiogalactoside), whereas binding site μ adopts State B (i.e., μ_o^B) displaying relatively high affinity toward Class G^b galactosides but comparatively low affinity for Class G^a galactosides; further, each m_o^A : μ_o^B dimer contains one thiol group whose reaction with N-ethylmaleimide inactivates the transporter unless blocked by galactoside binding at site m_o^A, while the second homologous thiol of the dimer is unreactive toward thiol reagents. Translocation of the m_o^A : μ_o^B dimer involves closing of g_o followed by opening of g_i, and causes the two thiols (as well as sites m and μ) to interchange roles in a symmetrical fashion: m_o^A : μ_o^B ↔ m_i^B : μ_i^A. In the presence of a substantial (negative) transmembrane Δμ~_H⁺, the m : μ dimer is postulated to undergo an electrogenic protein conformational change to a second form, ^*(m : μ), in which both sites m and μ possess low affinity toward internal Class G^b substrates; galactoside transport in both m : μ and ^*(m : μ) is assumed to be coupled to H⁺-symport (⁻OH-antiport) with a stoichiometry of approximately 1 : 1. Finally, five characteristic predictions of the half-sites model are outlined for further tests of its validity.     

Growing up under generalized violence: An ecological study of homicide rates and secular trends in age at menarche in Colombia, 1940s–1980s

We examined secular changes in mean age at menarche among 5577 Colombian women born between 1941 and 1989, and correlated those changes with nation-wide rates of homicide and real gross domestic product per capita (GDP) at the year of birth and at the year at age 5, within predefined historical periods. The mean (standard error) rate of change in age at menarche by year of birth was −0.55 (0.02) years/decade. The rate of change was not constant, but varied between historical periods as follows: −1.44, −0.14, −0.60, and −0.36 years/decade for the periods 1941–1947, 1948–1958, 1959–1978, and 1979–1989, respectively. The changes in age at menarche correlated positively with the changes in the nation-wide rates of homicide within such periods; i.e. decelerations in the menarcheal trend coincided with increases in the rates of homicide and vice versa. The correlation was higher with the rates of homicide when women were 5 years of age (r = 0.99, p = 0.01) compared to the rates of homicide at the year of birth (0.55, p = 0.45). There were negative correlations between the changes in age at menarche and the changes in GDP, but they were weaker than those with the rates of homicide. These results could suggest a potential impact on maturation of psychosocial stress in childhood due to exposure to a generalized atmosphere of violence and fear.     

Energy metabolism,body-temperature and breathing parameters in nontorpid blue-naped mousebirdsUrocolius macrourus

Summary Breathing frequencyF _r of resting blue-naped mousebirdsUrocolius macrourus lies between 50–70 per min and correlates directly with ambient temperatureT _a and energy metabolismM. The nocturnal mean energy intake per breath varies between 5.6–17.7 mJ/g. At highT _a the birds show gular fluttering with a relatively constantF _r of about 460 min^–1.M shows a constant absolute day-night difference of 25 J/g·h; the relative differences areT _a-dependent between 36–168% (lower values at lowerT _a). Thermal conductance is 2.10–2.15 J/g·h·°C (predicted 2.67), indicating a good insulation. Basal metabolic rate BMR is reduced by 63% compared to predicted values. At aT _a-range of +8–36 °C the birds are normothermic. Below this range nocturnalT _b andM decrease slightly with fallingT _a. The birds show partial heterothermia (shallow hypothermia). Clustering is an effective energy saving strategy which allows loweringM with keeping highT _b even at lowT _a.Oxygen-intake is controlled byF _r as well as by tidal volumeV t inT _a-dependent changing portions.V T can vary between 0.29–0.91 ml (mean value 49.7 ml).Abbreviations T _a ambient temperature - T _b body temperature - M energy metabolism - F _r breathing frequency - V T tidal volume - BMR basal metabolic rate - TNP thermoneutral point     

Effects of mine acid on the longevity and reproductive rate of the gastrotricha Lepidodermella squammata (Dujardin)

Effects of mine acid on longevity and reproductive rate of the parthenogenic gastrotrich Lepidodermella squammata were studied under laboratory culture conditions. Water from unpolluted and polluted streams was used, directly or mixed, to establish a series of test conditions at pH 8.1, 7.1, 6.4, 5.2, 4.6 and 3.3. Eggs, cultured individually under each test condition, were observed at 12 h intervals for hatching, daughter egg laying and death. Data representing 50 animals under each test condition were used in the construction of a series of life tables. From these were calculated maximal life expectancy (e_x), net reproductive rate per individual lifetime (R_o) and intrinsic rate of natural increase (r_max).Values of e_x, R_o and r_max were maximal at pH 7.1 and were reduced slightly at the higher pH; e_x was greatly reduced and R_o zero at pH 6.4 and 5.2; and e_x was zero at pH 4.6 and 3.3. Analysis of variance tests indicate significant differences between e_x values of L. squammata cultured at pH 8.1 and those cultured at pH 7.1, but no differences between R_o or r_max.Associated with the decrease in pH was an increase in total conductivity and a decrease in carbonate alkalinity and hence in carbonate conductivity. It appears that L. squammata is capable of living and reproducing at pH 6.0 to 6.5 under field conditions low in carbonates, providing non-carbonate ions are not abundant, or under field conditions high in non-carbonate ions, providing sufficient carbonates are present.     

Normal acid behavior for C(α)-proton transfer from a thiazolium lon

Rate constants for C(α)-proton transfer from racemic 2-(1-hydroxyethyl)-3,4-dimethylthi-oazolium ion catalyzed by lyoxide ion and various oxygen-containing and amine buffers were determined by iodination at 25°C and ionic strength 1.0 in H₂O. Thermodynamically unfavorable C(α)-proton transfer to oxygen-containing and amine bases shows general base catalysis with a Brønsted β value of ≥0.92 for bases of pK_a^′ ≤ 15; this indicates that the thermodynamically favorable protonation reaction in the reverse direction has a Brønsted α value ≤0.08, which is consistent with diffusion-controlled reprotonation of the C(α)-enamine by most acids. General base catalysis is detectable because there is an 85-fold negative deviation from the Brønsted correlation by hydroxide ion. Primary kinetic isotope effects of (k_H/k_D)_obsd = 1.0 for thermodynamically unfavorable proton transfer to buffer bases and hydroxide ion (ΔpK_a ≤ −6) and a secondary solvent isotope effect of k_DO⁻/k_HO⁻ = 2.3 for C(α)-proton transfer are consistent with a very late, enamine-like transition state and rate-limiting diffusional separation of buffer acids from the C(α)-enamine in the rate-limiting step, as expected for a “normal” acid. The second-order rate constants for catalysis by buffer bases were used to calculate a pK_a^′ of 21.8 for the C(α)-proton assuming a rate constant of 3 × 10^{9 −1} s⁻¹ for the diffusion-controlled reprotonation of the C(α)-enamine by buffer acids in the reverse direction. It is concluded (i) that C(α)-proton removal occurs at the maximum possible rate for a given equilibrium constant, and (ii) that C(α)-enamines can have a significant lifetime in aqueous solution and on thiamin diphosphate-dependent enzymes.     

A stochastic computer model for simulating population growth

A model is described for investigating the interactions of age-specific birth and death rates, age distribution and density-governing factors determining the growth form of single-species populations. It employs Monte Carlo techniques to simulate the births and deaths of individuals while density-governing factors are represented by simple algebraic equations relating survival and fecundity to population density. In all respects the model's behavior agrees with the results of more conventional mathematical approaches, including the logistic model andLotka's Law, which predicts a relationship betwen age-specific rates, rate of increase and age distribution. Situations involving exponential growth, three different age-independent density functions affecting survival, three affecting fecundity and their nine combinations were tested. The one function meeting the assumptions of the logistic model produced a logistic growth curve embodying the correct values or r_m and K. The others generated sigmoid curves to which arbitrary logistic curves could be fitted with varying success. Because of populational time lags, two of the functions affecting fecundity produced overshoots and damped oscillations during the initial approach to the steady state. The general behavior of age-dependent density functions is briefly explored and a complex example is described that produces population fluctuations by an egg cannibalism mechanism similar to that found in the flour beetle Tribolium. The model is free of inherent time lags found in other discrete time models yet these may be easily introduced. Because it manipulates separate individuals, the model may be combined readily with the Monte Carlo simulation models of population genetics to study eco-genetic phenomena.     

Transition-state structure for a conformation change of ribonuclease

The rate constants k₁₂ⁿ for isomerization of the E₁H isomer (pK_a 8 in H₂O) of ribonuclease-A to the E₂H isomer (pK_a = 6.1 in H₂O), determined from proton-uptake measurements by the temperature-jump technique, in mixtures of protium and deuterium oxides (atom fraction of deuterium n), are described by the equation k₁₂ⁿ = (733 ± 16)(1 − n + [0.46 ± 0.04]n)(1 − n + 0.69n)²sec⁻¹ at 25°C. On the basis of the absolute magnitude of the rate constant, the magnitude of the solvent isotope effect and the proton inventory, it appears that the rate-determining step is proton transfer to a water molecule from the imidazolium form of a histidine residue, with a product-like activated complex resembling a hydronium ion. The subsequent motion of the protein structure to generate the new isomer (conformation change) must then occur in a time approaching a vibrational period. Alternative but less likely mechanisms include rate-limiting protein reorganization concerted with proton transfer to water, rate-limiting diffusion of hydronium ion away from the enzyme, or “solvation catalysis” of protein reorganization.     

Dynamics of an age‐structured population drawn from a random numbers table

I constructed age‐structured populations by drawing numbers from a random numbers table, the constraints being that within a cohort each number be smaller than the preceding number (indicating that some individuals died between one year and the next) and that the first two‐digit number following 00 or 01 ending one cohort’s life be the number born into the next cohort. Populations constructed in this way showed prolonged existence with total population numbers fluctuating about a mean size and with long‐term growth rate (r) ≈ 0. The populations’ birth rates and growth rates and the females’ per capita fecundity decreased significantly with population size, whereas the death rates showed no significant relationship to population size. These results indicate that age‐structured populations can persist for long periods of time with long‐term growth rates of zero in the absence of negative‐feedback loops between a population’s present or prior density and its birth rate, growth rate, and fecundity, contrary to the assumption of density‐dependent regulation hypotheses. Thus, a long‐term growth rate of zero found in natural populations need not indicate that a population’s numbers are regulated by density‐dependent factors.     

Protein production from whey usingPenicillium cyclopium; growth parameters and cellular composition

Summary The growth parameters ofPenicillium cyclopium have been evaluated in a continuous culture system for the production of fungal protein from whey. Dilution rates varied from 0.05 to 0.20 h^–1 under constant conditions of temperature (28°C) and pH (3.5). The saturation coefficients in the Monod equation were 0.74 g l^–1 for lactose and 0.14 mg l^–1 for oxygen, respectively. For a wide range of dilution rates, the yield was 0.68 g g^–1 biomass per lactose and the maintenance coefficient 0.005 g g^–1 h^–1 lactose per biomass, respectively. The maximum biomass productivity achieved was 2 g l^–1 h^–1 biomass at dilution rates of 0.16–0.17 h^–1 with a lactose concentration of 20 g l^–1 in the feed. The crude protein and total nucleic acid contents increased with a dilution rate, crude protein content varied from 43% to 54% and total nucleic acids from 6 to 9% in the range of dilution rates from 0.05 to 0.2 h^–1, while the Lowry protein content was almost constant at approximately 37.5% of dry matter.Nomenclature (mg l^–1) C_o initial concentration of dissolved oxygen - (h^–1) D dilution rate - (mg l^–1) K₀₂ saturation coefficient for oxygen - (g l^–1) K_s saturation coefficient for substrate - (g g^–1 h^–1) lactose per biomass) m maintenance energy coefficient - (mM g^–1 h^–1O₂ per biomass) Q₀₂ specific oxygen uptake rate - (g l^–1) S residual substrate concentration at steady state - (g l^–1) S_o initial substrate concentration in feed - (min) t_1/2 time when C_o is equal to C_o/2 - (g l^–1) X biomass concentration - (g l^–1) X biomass concentration at steady state - (g g^–1 biomass per lactose) Y_G yield coefficient for cell growth - (g g^–1 biomass per lactose) Y_x/s overall yield coefficient - (h^–1) specific growth rate     

Backward bifurcation,equilibrium and stability phenomena in a three-stage extended BRSV epidemic model

In this paper we consider the phenomenon of backward bifurcation in epidemic modelling illustrated by an extended model for Bovine Respiratory Syncytial Virus (BRSV) amongst cattle. In its simplest form, backward bifurcation in epidemic models usually implies the existence of two subcritical endemic equilibria for R ₀ < 1, where R ₀ is the basic reproductive number, and a unique supercritical endemic equilibrium for R ₀ > 1. In our three-stage extended model we find that more complex bifurcation diagrams are possible. The paper starts with a review of some of the previous work on backward bifurcation then describes our three-stage model. We give equilibrium and stability results, and also provide some biological motivation for the model being studied. It is shown that backward bifurcation can occur in the three-stage model for small b, where b is the common per capita birth and death rate. We are able to classify the possible bifurcation diagrams. Some realistic numerical examples are discussed at the end of the paper, both for b small and for larger values of b.      

On the average cellular volume in synchronized cell populations

As was done by Sinclair and Ross (1969(, we consider a cellular population that consists initially (at time zero) ofN ₀ newborn cells, all with the same volumev _o. It is assumed that the occurrence of cell division is determined only by a cell’s age, and not by its volume. The frequency function of interdivision times, τ, is denoted byf(τ). If cell death is negligible, the expected number of cells,N(t), will increase according to the laws of a simple age-dependent branching process. The expression forN(t) is obtained as a sum over all generations; thevth term of this sum, in turn, is a multiple convolution integral, reflecting the life history ofvth generation cells (i.e., the lengths of thev successive interdivision periods plus the age of the cell at timet). Assuming that cell volume is a given function of cell age, e.g., linear or exponential, and that cellular volume is exactly halved at each division, it is possible to calculate the volume of a cell with a given life history, and thus the average cellular volume of the whole population as a function of time. If at time zero the volumes differ from cell to cell, the final equation must be modified by averaging over initial volumes. In the case of linear volume increase with age, a very simple asymptotic expression is found for the average cellular volume ast→∞. The case of exponential volume increase with age also leads to a simple asymptotic formula, but the resulting volume distribution is unstable. The mean cellular volume at birth and the second moment of the volume distribution can be calculated in a similar manner. Work supported by the U.S. Atomic Energy Commission.     

Leaf Longevity,Construction, and Maintenance Costs of Three Mangrove Species Under Field Conditions

This study assessed the effect of leaf age on construction cost (CC) in the mangrove species Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle growing in their natural habitat. Leaf osmolality values were species-specific, the highest in A. germinans (1 693 mmol kg^–1) and the lowest in L. racemosa (1 270 mmol kg^–1). In the three species, contents of chlorophyll (a+b) (Chl_a+b) and nitrogen (N) per unit of leaf area were maximal in adult leaves and tended to decline with age. Leaf mass to leaf area ratio (LMA) and ash content increased during leaf ageing. Similarly, as leaves aged, a significant increase in leaf construction cost per leaf area (CC_a) was observed, while per leaf mass (CC_m) it remained almost constant, suggesting a sustained production of leaf compounds as leaves became older. CC was positively correlated with LMA and heat of combustion (Hc) per leaf area, suggesting differences among species in the quantity and composition of expensive compounds. Leaf half lifetime (t_0.5) showed contrasting values in the three mangrove species (60, 111, and 160 d in L. racemosa, R. mangle, and A. germinans, respectively). Overall, L. racemosa was the species with less expensive leaves to construct while leaves of A. germinans and R. mangle had the highest CC_m and CC_a, respectively. Leaf longevity was positively correlated with the ratio between CC and maximum photosynthetic rate (P _max), clearly showing the existence of a balance between leaf costs and benefits.