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1 Downloaded from UvA-DARE, the institutional repository of the University of Amsterdam (UvA) File ID Filename Version uvapub:20101 Chapter 4 Inferring demographic processes from population size structure in corals unknown SOURCE (OR PART OF THE FOLLOWING SOURCE): Type PhD thesis Title Evolutionary Ecology of the coral genus Madracis - an illustration of the nature of species in scleractinian corals Author(s) M.J.A. Vermeij Faculty FNWI: Institute for Biodiversity and Ecosystem Dynamics (IBED), FNWI: Institute for Biodiversity and Ecosystem Dynamics (IBED) Year 2002 FULL BIBLIOGRAPHIC DETAILS: Copyright It is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), other than for strictly personal, individual use, unless the work is under an open content licence (like Creative Commons). UvA-DARE is a service provided by the library of the University of Amsterdam ( (pagedate: )
2 Chapter 4 Inferring demographic processes from population size structure in corals Vermeij M.J.A. and R.P.M. Bak Chapter 4 Analyzing population size structure 61
3 ABSTRACT Analyzing the size structure in coral population is difficult because corals vary enormously in size. The great differences in colony size are caused by the coral colony growth process. We studied size distributions in coral species and found that analyzing coral size frequencies using log transformations of colony size has a number of advantages. Coral size distributions become comparable within and between species across space and through time. Such analyses indicate that (1) small colony size and positively skewed size-frequency distributions characterize populations of brooding species. (2) Spawning species show large colony size and negatively skewed size-frequency distributions. (3) Populations in marginal habitats are characterized by a coefficient of variation higher than The described method of log transforming size frequency data of coral populations provides both scientists and wildlife managers with a new tool to monitor the dynamics of coral populations. INTRODUCTION Analysis of coral population size structure is difficult because corals vary enormously in size. Large variation in coral colony size occurs within populations of species such as Porites or Montastraea that have massive colonies reaching sizes of > cm 2. Great differences in colony size also hamper comparison between species, e.g. between small species such as Agaricia humilis or Madracis decactis (< 15 cm 2 ) and species such as Porites or Montastraea. Coral populations are dominated by small size classes (Soong 1993, Bak & Meesters 1998, Meesters et al. 2000) and it is essential to include these in analyses. Clearly a problem arises if one wishes to compare populations including <1 to cm 2 large colonies on a linear scale. The great differences in colony size are caused by the coral colony growth process. Corals grow through division of the first initial polyp, followed by division of subsequent polyps, as an exponential function' Power functions are however often used to describe the increase of corals as increasing semi circles, semi spheres or complications of such shapes. An underlying biological process, rather than the mathematical one, is however difficult to discover for power functions (x 2 ). Such growth leads to a potentially very rapid enlargement of coral living surfaces after the initial multiplication of the first polyps. Differences in coral longevity or colony partial mortality result in great intra and inter specific differences in maximum colony size. The enormous variation of colony size also implies that size differences, seen on linear scale, are not always very meaningful. Loss of 10 cm 2 has a different biological meaning for a small Chapter 4 Analyzing population size structure 62
4 Colony size (P; in cm 2 or no. of polyps) Figure 1. The relation between colony size and the number of corresponding iterations (t) using two different bases for the log transformation. Base= 2 ( ) and base= 10 (o). colony (e.g. 20 cm 2 ), than for a large one (e.g cm 2 ). Size is relatively easy to measure and an important character in populations of clonal organisms since it relates better than age to life history parameters such as mortality and reproductive output. We studied size distributions in coral species and found that analyzing coral size frequencies using log transformations of colony size has a number of advantages. Coral size distributions become comparable within and between species. Resolution of differences between species and in small size classes is much increased. Size A smaller base results in more iteration steps, i.e. a higher descriptive resolution, over frequencies tend to become approximately normal and can be characterized by the descriptors of such distributions: skewness, coefficient of variation etc. Here we want to explain the method of transformation and give some examples of applications to coral populations. METHODS Log transforming size data If we assume that a colony doubles in size at every iteration (i.e. growth step in time), a 10cm 2 increase in size for a juvenile coral of 0.5cm 2 corresponds to more growth iterations than for a lm 2 colony. To distinguish between the different implications of a 10cm 2 size increase for small and large colonies we prefer an indication of relative change in colony size. Relative change in size is then represented by the number of iterations that are required to bring about the increase in colony size. Since the change in size now depends on the number of iterations we Chapter 4 Analyzing population size structure 53
5 hypothesize that the number of iterations corresponding to an absolute size increase better describes relative differences between species than absolute size. The number of iterations or time steps (t), in case of undisturbed growth, can be calculated, assuming a theoretical colony whose polyps divide at eveiy growth step, using the exponential growth function: Pt = P 2' (1) Pt is the colony size (expressed in cm or as the number of polyps, P) after t growth intervals, i.e. iterations, in which the original colony size P 0 doubles every iteration. The division rate of polyps will eventually decrease, because of morphological and physiological constraints, and deviations from the exponential growth curve can be expected. The exponential function addresses the basic process in colony growth, i.e. polyp multiplication. The function representing real growth is probably an interaction of exponential growth functions and physiological or developmental constraints. k in *- Before transformation After transformation Figure 2. Size-frequency of Madracis pharensis colonies (n= 638). The untransformed (above) and log-transformed size-frequency (below) illustrate the difference in resolution in the small colony size classes. After the initial growth phase of a coral, a larger range along the x-axis corresponds to less iterations (Fig. 1) and a large range along the x-axis corresponds to a small range along the y-axis. It shows that the descriptive resolution in small sized colonies, i.e. range along the y-axis, Chapter 4 Analyzing population size structure 64
6 becomes higher for the first iteration steps. Larger sizes represent a relatively small number of iterations, a small part of the iterative process. To describe variation in colony size we need a method where colony size is replaced by a number of iterations. Logarithmic transformation of the x-axis, i.e. colony size, is by far the easiest way to obtain the number of iterations from the size data. We can visualize the distribution of colonies in a histogram where the size-categories represent successive growth steps based on the iteration steps mentioned above. The number of iterations determines the descriptive resolution. Figure 1 shows that the base of the log transformation (2 or 10, which were chosen for illustrative purposes and have no biologically relevance) does not affect the basic pattern. It only affects the number of iterations that fit in a given size-range along the x- axis, i.e. it affects the resolution. Therefore, the absolute size range of a species determines which base should be used. Increasing the number of iterations over the size range increases the resolution of the model. Smaller bases are used for species with a small size range (i.e. In2 for small species, < 400 cm maximum colony size) whereas large bases, i.e. In 10 should be used for larger species. The transformation is the following, where the number of iterations is the new approximation of colony size. o- L iteration //->\ Size = base (2) Iteration = In (size) /In (base) (3) A log transformation of size distribution data results in an approximately normal distribution, because it increases the number of smaller size classes while size classes containing larger colonies are reduced. In general, high juvenile mortality depletes the number of colonies present in the smallest size-classes. The effect of whole colony mortality gradually decreases with increasing size (Barry and Tegner 1989) Partial mortality and fission limit the number of individuals in the larger sizeclasses (Bak and Meesters 1998, 1999). This results in an influx of colonies towards the intermediate size classes. Environmental variation and species-specific components result in deviations from this general shape. We analyzed the differences in transformed and untransformed size distributions of Madracis pharensis colonies (Fig. 2; n = 638) at different depths. The colonies were measured in a 150m 2 beltransect at Buoy 1, Curaçao, Netherlands Antilles (12 05'N, 69 00'W) over a 10 to 60m depth range. The normal distribution has mathematical properties that can be described using various statistics that are an useful tool in the in the study of coral size distributions. Statistics that have been used in Chapter 4 Analyzing population size structure 65
7 reproductive output t * mortality mortality mortality Figure 3. Theoretical life cycle of corals. The transitions, between the four hypothetical stages representing log transformed size classes, can be changed to create new size distributions. studies on coral populations (Bak and Meesters 1998, 1999, Meesters et al. 2000, Vermeij and Bak subm.) are the coefficient of variation (CV), skewness (gl), the mode (Mo), mean size after transformation (Ms) and' the standard deviation of transformed data (SD). These statistics (Bendel et al. 1989, Sokal and Rohlf 1981) describe the shape of a distribution and allow comparisons between species and environments independent of size. RESULTS AND DISCUSSION Statistics of size frequency distributions Input of juvenile corals In a population having no input of juveniles during most of the year, such as in spawning species, the first size classes become depleted in coral colonies (Fig. 4b). All juveniles grow into larger size classes while there is no recruitment. Continuous input of juveniles, as in some brooding species, results in a high number of juveniles in the smallest size classes (Fig. 4c). These contrasting reproductive modes can be characterized by the distribution statistics. Populations of brooding species are characterized by increased variation in size (CV) and are highly skewed to the right (positive gl). Juvenile mortality Increased juvenile mortality decreases the number of juveniles present in the first size classes causing a relative overrepresentation of large colonies in the population indicated by negative gl-values (Fig. 4d). Due the larger size range variation increases as well the mean colony size (Ms). Chapter 4 Analyzing population size structure 66
8 Partial mortality Partial mortality results in a lower number of colonies present in the largest size classes due increased transition possibilities to lower size classes (Fig. 4e). Colonies become over represented in intermediate size classes due to growth of colonies in lower size classes and due to partial mortality of colonies in higher size-classes. This results in lower over-all variation (CV) and a lower mean colony size (Ms). Environmental degradation Environmental degradation was modeled by combining the effects of increased juvenile and partial mortality and reducing transition probabilities into higher size classes, i.e. a slow down of growth by 30%. The juvenile fraction of the population is most affected and the relative high proportion of larger colonies causes negative gl-values and the variation to decrease proportionate to the mean (CV decreases). (A) skew 1,80 CV 0.69 mean 5.06 stdev 8.23 (D) skew CV 0.64 mean 1.62 stdev o T- CD m T LO T- ID K) CM -r- CD CO T CD ID CM CO CD CM (B) skew 0.95 (E) CV 0.87 mean 6.82 H ^ _ stdev 8.15 skew 4.99 CD CM T- ^r en T- co m CM T- CD CO TT =1- CD CO CM CD CD (C) skew 7.36 CV 0.21 mean 2.55 stdev (F) skew CV 0.42 mean stdev Figure 4. The modeled effects of change in demographic processes in a coral population, (a) Model input data representing a natural size distribution, (b) no juvenile input, i.e. reproductive output is zero, representing a spawning coral species outside the reproductive season, (c) increased input of juveniles (reproductive output 10 times enhanced), (d) increased juvenile mortality (10 times enhanced), (e) increased partial mortality (probability for transitions to lower size class increase 3 times in size-classes above the mode), (f) environmental degradation simulated by increasing juvenile mortality (x2), increasing the transition to lower size classes and reducing growth, i.e. reducing transition probabilities to higher size classes by 70%. All transition values are hypothetical. Chapter 4 Analyzing population size structure 67
9 3 P 2 ai ös 1 c S OT O v* % -1 y=-0.426ln(x) R 2 = 0.41 o 8 a Colony size (cm 2 ) Figure 5. The relation between skewness (gl) and mean colony size indicates that brooding species ( ) are characterized by small colony size and positively skewed populations. This indicates that juveniles dominate these populations. Spawning species (o) have a large mean size and their populations are more negatively skewed. Data are size frequencies of 19 Caribbean coral species (Meesters et al. 2000, Vermeij and Bak subm.) > u * *»^\ o O 0.4 o O 0.3 <b y = Ln(x) = 0.61 u.u Abundance (n / m 2 ) O Figure 6. The relation between the coefficient of variation (CV) and abundance (n m" ) for six Madracis species. Low abundance of colonies in populations at the margins of a species vertical distribution ( ) corresponds to higher values for CV (> 0.50). Populations in non-marginal habitats are indicated by (o). Chapter 4 Analyzing population size structure 68
10 The model shows how the statistics can be indicative of the processes that changed the initial size distributions and illustrates their potential in characterizing species life history strategies (Fig. 4 b-e) or recognizing populations growing in degrading habitats (Fig. 4f). Comparing statistics between species and populations The use of descriptors is not limited to comparing size frequencies; they can be used to find general ecological patterns in coral species on reefs. The relation between mean colony size and skewness is shown for four populations of 19 different species, 7 gamete spawning and 12 brooding species in Fig. 5 (see Meesters et al. 2001, Vermeij and Bak subm.). It appears that small colony size and positively skewed size distributions characterize populations of brooding species. This must be due to the relatively high input over longer time periods of recruits compared to spawning species. These differences correspond to the differences detected in the two modes of reproduction modeled in Fig. 4b and c. We suggest that the relation between CV and abundance (Fig. 6) can be used to characterize populations in sub-optimal or marginal habitats for six Caribbean Madracis species (data Vermeij and Bak subm.). Madracis species were investigated over a depth gradient from 10 till 50m within isobathic 30m2 belt transects at each 10m depth. At the upper and lower part of species depth range colonies were less abundant and these depths must be considered as marginal habitats. There is a significant relation between CV and abundance (p < 0.000; Fig. 6) and for these Madracis populations it appears that a CV > 0.50 characterizes populations in suboptimal conditions. IN CONCLUSION In this paper we have shown the method and advantage of logtransforming coral size data for analytical purposes, such as the characterization of patterns in life history strategies or recognizing populations in marginal habitats. Using a matrix-model the expected effects of changes in life history processes were tested for their effect on the coral size distributions. The effects are expressed in the statistics skewness (gl), Coefficient of Variation (CV), mean size after logtransformation (Ms). The use of log-transformed size frequencies indicates the practical use of these statistics for population ecologists and wildlife managers interested in understanding the dynamics of coral populations. Chapter 4 Analyzing population size structure 69
11 CITED LITERATURE Bak RPM, Meesters EH (1998) Coral population structure: the hidden information of colony size-frequency distributions. Mar Ecol Prog Ser 162: Bak RPM, Meesters EH (1999) Population structure as a response of coral communities to global change. Am Zool 39(1): Barry JP, Tegncr MJ (1983) Inferring demographic processes from size-frequency distributions: Simple models indicate specific patterns of growth and mortality. Fishery Bull U.S. 88: Bendel RB, Higgins SS, Teberg JE, Pyke PA (1989) Comparison of skewness coefficient, coefficient of variation and Gini coefficient as inequality measures within populations. Oecologia 78: Caswell H (1989) Matrix population models: construction, analysis, and interpretation. Sinauer Associates Inc., Sunderland, USA. Meesters EH, Hilterman M, Kardinaal E, Keetman M, de Vries M, Bak, RPM (2000) Colony size-frequency distributions of scleractinian populations: Spatial and interspecific variation. Mar Ecol Prog Ser 209: Sokal RR, Rohlf FJ (1981) Biometry. W. H. Freeman and Company, New York USA. Soong K (1993) Colony size as a species characteristic in massive reef corals. Coral Reefs 12(2): Vermeij MJA, Bak RPM (submitted) Species specific population structure of closely related coral morphospecies along a depth gradient (5-60m) over a Caribbean reef slope. Bull Mar Sc. Wells JW (1973a) New and old corals from Jamaica. Bull Mar Sc 23(1): Wells JW (1973b) Two new hermatypic corals from the West Indies. Bull Mar Sc 23(4): Acknowledgements. We thank the Carmabi Foundation for logistical support. Wc are grateful for comments and discussions with Dr Erik Meesters, which have inspired and improved the manuscript. An anonymous reviewer suggested useful improvements. Chapter 4 Analyzing population size structure 70
Downloaded from UvA-DARE, the institutional repository of the University of Amsterdam (UvA) http://hdl.handle.net/11245/2.122992
Downloaded from UvA-DARE, the institutional repository of the University of Amsterdam (UvA) http://hdl.handle.net/11245/2.122992 File ID Filename Version uvapub:122992 1: Introduction unknown SOURCE (OR
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