More than the sum of the parts: annual partitioning within spatial guilds underpins community regulation.

To withstand the pressures of a rapidly changing world, resilient ecosystems should exhibit compensatory dynamics, including uncorrelated temporal shifts in population sizes. The observation that diversity is maintained through time in many systems is evidence that communities are indeed regulated and stabilized, yet empirical observations suggest that positive covariance in species abundances is widespread. This paradox could be resolved if communities are composed of a number of ecologically relevant sub-units in which the members compete for resources, but whose abundances fluctuate independently. Such modular organization could explain community regulation, even when the community as a whole appears synchronized. To test this hypothesis, we quantified temporal synchronicity in annual population abundances within spatial guilds in an estuarine fish assemblage that has been monitored for 36 years. We detected independent fluctuations in annual abundances within guilds. By contrast, the assemblage as a whole exhibited temporal synchronicity-an outcome linked to the dynamics of guild dominants, which were synchronized with each other. These findings underline the importance of modularity in explaining community regulation and highlight the need to protect assemblage composition and structure as well as species richness.

Relationship between evenness and body size in species rich assemblages.

Evenness is a key measure of community structure. Here, we examine the relationship between evenness and size-abundance distributions for both individuals and species using data gathered from Amazonian fish assemblages. We show that evenness increases as the fraction of numerically abundant species in larger body-size classes rises. As any processes that enable larger bodied species to increase their numerical dominance will influence evenness, these results help explain why evenness is an important correlate of ecosystem function.

Local and regional rarity in a diverse tropical fish assemblage.

Because most species in an ecological assemblage are rare, much of the species richness we value is due to taxa with few individuals or a restricted distribution. It has been apparent since the time of ecological pioneers such as Bates and Darwin that tropical systems have disproportionately large numbers of rare species, yet the distribution and abundance patterns of these species remain largely unknown. Here, we examine the diversity of freshwater fish in a series of lakes in the Amazonian várzea, and relate relative abundance, both as numbers of individuals and as biomass, to the occurrence of species in space and time. We find a bimodal relationship of occurrence that distinguishes temporally and spatially persistent species from those that are infrequent in both space and time. Logistic regression reveals that information on occurrence helps distinguish those species that are rare in this locality but abundant elsewhere, from those that are rare throughout the region. These results form a link between different approaches used to evaluate commonness and rarity. In doing so, they provide a tool for identifying species of high conservation priority in poorly documented but species rich localities.

Biological diversity of fish communities: pattern and process.

For over 150 years, ecologists have been striving to explain fundamental patterns of biological diversity, such as the observation that communities invariably consist of common and rare species, and to unravel the processes that underpin these patterns. This task is increasingly urgent given the accelerating loss of biological diversity. Although fishes are the most diverse vertebrate taxon and fish communities occur in a wide range of habitats, they have been relatively little studied in the quest to elucidate the processes that shape patterns of biological diversity. Here, some of the topics that investigations of fish assemblages can illuminate are highlighted. These include the characteristics of ecological communities and the role that dispersal limitation plays in structuring them, the distinction between core and occasional species, the insights that evaluating abundance in different currencies can bring and the assessment of community capacity. Questions are identified that future investigations of fish communities might tackle and a case study of a biodiverse ecoregion (Thailand and Peninsula Malaysia) is used to illustrate the need for better links between these ecological questions and effective conservation practice.

Effects of polyandry on male phenotypic diversity.

Polyandry has the potential to affect the distribution of phenotypes and to shape the direction of sexual selection. Here, we explore this potential using Trinidadian guppies as a model system and ask whether polyandry leads to directional and/or diversifying selection of male phenotypic traits. In this study, we compare the phenotypic diversity of offspring from multiply and singly sired broods. To quantify phenotypic diversity, we first combine phenotypic traits using multivariate methods, and then take the dispersion of individuals in multivariate space as our measure of diversity. We show that, when each trait is examined separately, polyandry generates offspring with a higher proportion of bright coloration, indicating directional selection. However, our multivariate approach reveals that this directionality is accompanied by an increase in phenotypic diversity. These results suggest that polyandry (i) selects for the production of sons with the preferred brighter colour phenotypes whereas (ii) enhancing the diversity of male sexual traits. Promoting phenotypic diversity may be advantageous in coping with environmental and reproductive variability by increasing long-term fitness.

The effect of social environment during ontogeny on life history expression in the guppy Poecilia reticulata.

The effects of the social environment during development on life-history decisions and adult behaviour were assessed using male guppies Poecilia reticulata. Males raised with adults developed secondary sexual characteristics later than males raised either singly or with four of their siblings indicating social inhibition of maturation was evident in P. reticulata. There was no effect, however, of rearing environment on male behaviour. The results reveal that social environment during development can influence life-history decisions but is less important than immediate social context in determining male behavioural phenotype in P. reticulata.

Sex and differentiation: population genetic divergence and sexual dimorphism in Mexican goodeid fish.

Genetic differentiation arises due to the interaction between natural and sexual selection, migration and genetic drift. A potential role of sexual selection in speciation has received much interest, although comparative studies are inconsistent in finding supporting evidence. A poorly tested prediction is that species subject to a higher intensity of sexual selection should show greater genetic differentiation amongst populations because females from these populations should be more choosy in mate choice. The Goodeinae is a group of endemic Mexican fishes in which female choice has driven some species to be morphologically sexually dimorphic, whereas others are relatively monomorphic. Here, we measured population divergence, using microsatellite loci, within four goodeid species which show contrasting levels of sexual dimorphism. We found higher levels of differentiation between populations of the more dimorphic species, implying less gene flow between populations. We also found evidence of higher levels of genetic differences between the sexes within populations of the dimorphic species, consistent with greater dispersal in males. Adjusted for geographic distance, the mean F(ST) for the dimorphic species is 0.25 compared with 0.16 for the less dimorphic species. We conclude that population differentiation is accelerated in more sexually dimorphic species, and that comparative phylogeography may provide a more powerful approach to detecting processes, such as an influence of sexual selection on differentiation, than broad-scale comparative studies.

Gametic isolation in guppies (Poecilia reticulata)

Post-mating reproductive isolating mechanisms may be among the earliest reproductive barriers to emerge among incipient species. Trinidadian guppy, Poecilia reticulata, populations in the Caroni and Oropouche drainages in Northern Trinidad exhibit marked genetic divergence and provide an ideal system in which to search for these barriers. We inseminated virgin females with equal amounts of sperm from two males, a 'native' male from the female's own population and a 'foreign' male from the other drainage. Artificial insemination ensured that mating order and mate choice did not affect the outcome. Paternities were assigned to the resulting broods using microsatellite markers. As predicted, sperm from native males had precedence over foreign sperm. Moreover, this effect was symmetrical for both drainages. In contrast, we detected no native sperm precedence in controls, in which females received sperm from the same and another population within the same drainage. Our results show that gametic isolation can arise between geographically proximate, though genetically divergent, populations of a single species and highlight the potential role of this process in speciation.

Gametic isolation in guppies (Poecilia reticulata).

Post-mating reproductive isolating mechanisms may be among the earliest reproductive barriers to emerge among incipient species. Trinidadian guppy, Poecilia reticulata, populations in the Caroni and Oropouche drainages in Northern Trinidad exhibit marked genetic divergence and provide an ideal system in which to search for these barriers. We inseminated virgin females with equal amounts of sperm from two males, a 'native' male from the female's own population and a 'foreign' male from the other drainage. Artificial insemination ensured that mating order and mate choice did not affect the outcome. Paternities were assigned to the resulting broods using microsatellite markers. As predicted, sperm from native males had precedence over foreign sperm. Moreover, this effect was symmetrical for both drainages. In contrast, we detected no native sperm precedence in controls, in which females received sperm from the same and another population within the same drainage. Our results show that gametic isolation can arise between geographically proximate, though genetically divergent, populations of a single species and highlight the potential role of this process in speciation.

Intrinsic reproductive isolation between Trinidadian populations of the guppy, Poecilia reticulata.

Although Trinidadian populations of the guppy, Poecilia reticulata, show considerable adaptive genetic differentiation, they have been assumed to show little or no reproductive isolation. We tested this assumption by crossing Caroni (Tacarigua River) and Oropuche (Oropuche R.) drainage populations from Trinidad's Northern Range, and by examining multiple aspects of reproductive compatibility in the F1, F2 and BC1 generations. In open-aquarium experiments, F1 males performed fewer numbers of mating behaviours relative to parental population controls. This is the first documentation of hybrid behavioural sterility within a species, and it suggests that such sterility may feasibly be involved in causing speciation. The crosses also uncovered hybrid breakdown for embryo viability, brood size and sperm counts. In contrast, no reductions in female fertility were detected, indicating that guppies obey Haldane's rule for sterility. Intrinsic isolation currently presents a much stronger obstacle to gene flow than behavioural isolation, and our results indicate that Trinidadian populations constitute a useful model for investigating incipient speciation.

Patterns of speciation in endemic Mexican Goodeid fish: sexual conflict or early radiation?

Currently there is much interest in the potential for sexual selection or conflict to drive speciation. Theory proposes that speciation will be accelerated where sexual conflict is strong, particularly if females are ahead because mate choice will accentuate divergence by limiting gene flow. The Goodeinae are a monophyletic group of endemic Mexican fishes with an origin at least as old as the Miocene. Sexual selection is important in the Goodeinae and there is substantial interspecific variability in body morphology, which influences mate choice, allowing inference of the importance of female mate choice. We therefore used this group to test the relationship between sexual dimorphism and speciation rate. We quantified interspecific variation in sexual dimorphism amongst 25 species using a multivariate measure of total morphological differentiation between the sexes that accurately reflects sexual dimorphism driven by female mate choice and also used a mtDNA-based phylogeny to examine speciation rates. Comparative analyses failed to support a significant association between sexual dimorphism and speciation rate. In addition, variation in the time course of speciation throughout the whole clade was also examined using a similar tree containing 34 extant species. A constant rates model for the growth of this clade was rejected, but analyses instead indicated a decline in the rate of speciation over time. These results support the hypothesis of an early expansion of the group, perhaps due to an early radiation influenced by the key innovation of live bearing, or the prevalence of Miocene volcanism. In general, support for the role of sexual selection in generating patterns of speciation is proving equivocal and we argue that vicariance biogeography and adaptive radiations remain the most likely determinants of major patterns of diversification of continental organisms.

Monitoring change in biodiversity through composite indices.

The need to monitor trends in biodiversity raises many technical issues. What are the features of a good biodiversity index? How should trends in abundance of individual species be estimated? How should composite indices, possibly spanning very diverse taxa, be formed? At what spatial scale should composite indices be applied? How might change-points--points at which the underlying trend changes--be identified? We address some of the technical issues underlying composite indices, including survey design, weighting of the constituent indices, identification of change-points and estimation of spatially varying time trends. We suggest some criteria that biodiversity measures for use in monitoring surveys should satisfy, and we discuss the problems of implementing rigorous methods. We illustrate the properties of different composite indices using UK farmland bird data. We conclude that no single index can capture all aspects of biodiversity change, but that a modified Shannon index and the geometric mean of relative abundance have useful properties.

Multiple mating and reproductive skew in Trinidadian guppies.

Male offspring production in promiscuously mating species is typically more skewed than female offspring production. It is therefore advantageous for males to seek as many mating partners as possible. However, given the documented benefits of polyandry we expect females, as well as males, to mate multiply. We tested these ideas using Trinidadian guppies, Poecilia reticulata. Fishes were collected from the wild, housed in groups of 10 males and 10 females and allowed to reproduce freely over a period of three months. We used hypervariable microsatellite loci to identify the parents of 840 offspring and to quantify the variance in mating success. As anticipated, and in line with the Bateman gradient, there was greater skew in the number of progeny produced by males. By contrast, we found no sex difference in mating partner number over the duration of the experiment. A median of two males fathered each brood and there was marked turnover in the identities of the sires of successive broods. Female partner turnover was, however, less than expected under random mating. We suggest that partner switching over time, as well as polyandry within broods, could contribute to the maintenance of genetic diversity in guppy populations.

Multiple mating and reproductive skew in Trinidadian guppies

Male offspring production in promiscuously mating species is typically more skewed than female offspring production. It is therefore advantageous for males to seek as many mating partners as possible. However, given the documented benefits of polyandry we expect females, as well as males, to mate multiply. We tested these ideas using Trinidadian guppies, Poecilia reticulata. Fishes were collected from the wild, housed in groups of 10 males and 10 females and allowed to reproduce freely over a period of three months. We used hypervariable microsatellite loci to identify the parents of 840 offspring and to quantify the variance in mating success. As anticipated, and in line with the Bateman gradient, there was greater skew in the number of progeny produced by males. By contrast, we found no sex difference in mating partner number over the duration of the experiment. A median of two males fathered each brood and there was marked turnover in the identities of the sires of successive broods. Female partner turnover was, however, less than expected under random mating. We suggest that partner switching over time, as well as polyandry within broods, could contribute to the maintenance of genetic diversity in guppy populations.

Inbreeding depression and genetic load of sexually selected traits: how the guppy lost its spots.

To date, few studies have investigated the effects of inbreeding on sexually selected traits, although inbreeding depression on such traits can play an important role in the evolution and ecology of wild populations. Sexually selected traits such as ornamentation and courtship behaviour may not be primary fitness characters, but selection and dominance coefficients of their mutations will resemble those of traits under natural selection. Strong directional selection, for instance, through female mate-choice, purges all but the most recessive deleterious mutations, and the remaining dominance variation will result in inbreeding depression once populations undergo bottlenecks. We analysed the effects of inbreeding on sexually selected traits (colour pattern and courtship behaviour) in the male guppy, Poecilia reticulata, from Trinidad, and found a significant decline in the frequency of mating behaviour and colour spots. Such effects occurred although the genetic basis of these traits, many of which are Y-linked and hemizygous, would be expected to leave relatively little scope for inbreeding depression. Findings suggest that these sexually selected traits could reflect the genetic condition or health of males, and thus may be informative mate-cue characters for female choice as suggested by the 'good genes' model.

Patterns of sperm precedence and predictors of paternity in the Trinidadian guppy.

Despite its widespread occurrence in animals, sperm competition has been studied in a limited range of taxa. Among the most neglected groups in this respect are internally fertilizing fish in which virtually nothing is known about the dynamics of sperm competition. In this study, we examined the outcome of sperm competition when virgin female guppies mated with two males. Behavioural cues were used to ensure that each male mated once (with female cooperation) and that sperm were successfully inseminated at copulation. Two polymorphic microsatellite loci were used to estimate the proportion of offspring sired by the second male (P2) and the results revealed a bimodal distribution with either first or (more often) second male priority The observed P2 distribution differed from that expected under the 'fair raffle' model of sperm competition. Random sperm mixing is therefore unlikely to account for the observed variance in P2 in this study. A further aim of our study was to identify predictors of male reproductive success. Using logistic linear modelling, we found that the best predictors of paternity were time to remating and the difference in courtship display rate between first and second males. Males that mated quickly and performed relatively high numbers of sigmoid displays obtained greater parentage than their slower and less vigorous counterparts. Since females are attracted to high-displaying males, our results suggest that female choice may facilitate sperm competition and/or sperm choice in guppies.

Sexual conflict and evolution in Trinidadian guppies.

Recent investigations have highlighted the importance of sexual conflict in the evolution of reproductive isolation. Examination of the Trinidadian guppy (Poecilia reticulata) shows how geographic variation in sexual conflict can mediate the emergence of isolating mechanisms. On the basis of pre-mating behaviour guppy females appear to be winning the battle of the sexes in low risk localities whereas males are apparently ahead in high-predation environments. However, the conclusion that sexual selection (through female choice) is replaced by sexual coercion of females (as a consequence of sneaky mating) in predator rich (and productive) assemblages takes no account of post-copulatory mechanisms. Recent work on sperm competition in guppies suggests coevolution between males and females may also occur in the post-mating, pre-zygotic arena. The potential for the evolution of reproductive isolation at each stage of the mating sequence is assessed.

Multiple benefits of multiple mating in guppies.

The rewards of promiscuity for males are undisputed. But why should a female mate promiscuously, particularly when her partners offer no resources other than sperm and increase her chances of succumbing to predation or disease? This question has been hotly debated but at present remains largely unresolved [Jennions, M. D. & Petrie, M. (2000) Biol. Rev. 75, 21-64]. One possibility is that females exploit postcopulatory mechanisms, such as sperm competition, to increase both the quality and quantity of their offspring. In this paper, we use the Trinidadian guppy, a species with a resource-free mating system, to test the hypothesis that females gain multiple benefits from multiple mating. Our results indicate that multiply mated females secure substantive advantages: They have shorter gestation times and larger broods, and they produce offspring with better developed schooling abilities and escape responses than their singly mated counterparts.

Measuring richness and evenness.

Surveying Natural Populations by L-A.C. Hayek and M.A. Buzas Columbia University Press, 1997. $69.00/£48.00 hbk, $28.00/£19.00 pbk (xvi+563 pages) ISBN 0 231 10240 2/0 231 10241 0.