Chaotic genetic patchiness in an intertidal limpet,Siphonaria sp.

Variation of 4 polymorphic enzymes was studied for 2 yr (1978 and 1979) in an undescribed species of Siphonaria, a pulmonate limpet, from a rocky shore at Rottnest Island, Western Australia. Depending on the locus, significant genetic differences were found among sites along 50 m of shore, between high and low portions of the shore within sites, between adults and recruits, and between recruits in the 2 yr. This genetic heterogeneity does not follow a simple, consistent pattern, but forms a shifting, ephemeral genetic patchiness best described as chaotic. This patchiness may result from temporal variation of numbers and genotypes of recruits, which leads to the proposal that planktonic dispersal, although causing uniformity on a large scale, can give rise to fine-scale genetic patchiness.     

Current shifts and kin aggregation explain genetic patchiness in fish recruits

The scales of population structure in marine species depend on the degree to which larvae from different populations are mixed in the plankton. There is an intriguing trend in marine population genetic studies of significant genetic structure for larvae, recruits, or populations at fine scales that is unpatterned across space and changes through time. This "chaotic genetic patchiness" suggests that larval pools are not well mixed in the plankton. However, few studies have been able to distinguish among potential causes of spatial and temporal genetic heterogeneity: changes in larval migration patterns, changes in environmental selection, or stochasticity caused by "sweepstakes" reproductive success of spawners creating detectable family structure. Here we use microsatellite markers to show that significant allele frequency shifts occurred sporadically in space and time for cohorts of recruits of Paralabrax clathratus (kelp bass) collected once every two weeks over two years from five sites in the Santa Barbara Channel, California, USA. We found that the pattern of genetic differentiation among cohorts was explained by a combination of (1) family structure in some cohorts, evidenced by half and full siblings, and (2) an indication of changes in larval delivery. It is unlikely but possible that environmental selection also plays a role. Although sampling of potential source populations was incomplete, cohorts arriving during western current flows show most genetic similarity with a population sample collected in the west, and cohorts arriving during current flows from the southeast show similarity with population samples collected in the south and east. Despite the family structure apparent in some cohorts, these "sweepstakes" events occur on too fine a scale to create lasting year class genetic structure. The results corroborate oceanographic models of larval dispersal, which suggest that larval mixing in the plankton is less extensive than previously believed.     

Local genetic structure and relatedness in a solitary mammal, Neotoma fuscipes

We used DNA microsatellites to investigate temporal and spatial patterns of local genetic differentiation and relatedness in a solitary mammal, the dusky-footed woodrat (Neotoma fuscipes). Patterns of genetic variation were measured relative to spatial clusters, or neighborhoods, of woodrats. We detected significant genetic differentiation among woodrat neighborhoods in two populations spanning multiple habitat types and densities. Estimates of θ _ST among neighborhoods ranged 0.034–0.075 and were comparable to levels reported in social mammals. Genetic differentiation at such a local scale is noteworthy because it occurred in the absence of any physical barriers to gene flow, suggesting that the patterns observed are linked to the nonrandom patterns of mating and dispersal that characterize woodrat social structure. Genetic differentiation and relatedness among neighborhoods were even higher when only resident females were analyzed. These results are consistent with a pattern of female philopatry and male-biased dispersal in woodrats. Geographic distance and relatedness were inversely correlated in adult females at intermediate densities, but not at low densities. Nonetheless, matrilineal genetic structure was apparent even at low woodrat densities based on estimates of θ _ST among neighborhoods of resident females that were significantly greater than zero and consistently greater than estimates including all individuals. In summary, this study demonstrates a matrilineal genetic structure in dusky-footed woodrats. In addition, our results support the idea that intermediate densities may be better at facilitating the formation of spatial kin clusters than either extreme. An erratum to this article can be found at     

High genetic variability and patchiness in a common Great Barrier Reef zoanthid (Palythoa caesia)

Allozyme electrophoretic analysis of seven polymorphic enzyme loci suggested that 1261 samples of Palythoa collected along 1765 km of the Great Barrier Reef during 1992–1993 were members of a single taxon, identified as Palythoa caesia Dana, 1846, with high external morphological variability and possibly encompassing several previously described species. Populations were slightly genetically differentiated (standardized genetic variance, F _ST=0.010, p<0.05), but there was no evidence of isolation by distance, or of the particular genetic distinction of geographic sets of reefs such as the Swains, as has been observed in other invertebrates. Differentiation was thought to be the result of random selection acting on patches of larvae, not the consequences of long-term reproductive isolation of populations.     

Quantifying site quality in a heterogeneous landscape: recruitment of a reef fish

Heterogeneity in site quality can play an important role in patterns of abundance and population dynamics. Yet, estimating site quality in natural systems can be problematic because site quality can (1) vary through ontogeny for a focal organism, leading to shifts in site quality with age, (2) be confounded with (or masked by) variation in traits of individuals populating the sites, and (3) be correlated with local density. For example, if high-quality sites attract more individuals but vital rates are density dependent, then observed vital rates will be relatively homogeneous in space despite strong heterogeneity in site quality. Here, we operationally define site quality for a reef fish as the mean survival time of juveniles transplanted to sites at a common density and size structure, with random assignment of individuals to sites to remove potential confounding effects of local variation in individual quality and density. Our assays using juvenile age classes of the six-bar wrasse (Thalassoma hardwicke) showed that site quality varied in space (i.e., among patch reefs) but was constant through time. Site quality increased with availability of the branching coral Pocillopora (which is used as a refuge), but decreased with density of a predator, the arc-eye hawkfish, Paracirrhites arcatus (which also uses Pocillopora). We experimentally added colonies of Pocillopora to reefs and (1) increased site quality, (2) enhanced natural settlement rates of six-bar wrasse, but (3) attracted more hawkfish predators, and (4) did not increase survival of juvenile fish under ambient densities. Our results suggest that Pocillopora increases site quality, but attracts greater densities of settlers and predators, resulting in increased density dependence and predation, which mask the underlying effects of Pocillopora on site quality (supporting the hypothesis of "cryptic density dependence"). Variation in site quality and the possible confounding effects of density and individual traits warrant more experimental study.     

Fine-scale patchiness and genetic heterogeneity of recruits of the corallivorous gastropod Drupella cornus

The size-frequency distributions and genetic composition of recruits of the corallivorous snail Drupella cornus (Röding, 1798) were examined in outbreak populations collected from Ningaloo Reef, Western Australia in August 1990. The recruits are found in groups on digitate Acropora spp. corals. Among coral colonies, mean lengths of recruits in our samples ranged from <9 to 22 mm, but the snails within a group were generally similar in size. Despite the fact that D. cornus has planktonic larvae, there were marked genetic differences between groups of recruits on different coral colonies. The relatively large genetic subdivision among groups of recruits within sites over distances <80 m was measured as a value of F _ST (standardized variance in allelic frequencies) of 0.044. This was three times the value from comparisons of pooled samples of recruits from areas up to 119 km apart, and six times as great at the genetic subdivision among populations of adults over a distance of 180 km. Much of the genetic heterogeneity among groups of recruits is associated with mean size of the snails. Taken together, the size-frequency distributions and the genetic differences indicate that recruits within the same coral colony shared a common history of settlement, suggesting a cohesiveness of groups of larvae. Although the mechanisms for this patchiness are not understood, one implication is that studies of size-frequency distributions and genetic composition of cohorts of D. cornus must treat the group, not the individual snail, as the unit of replication.     

Effects of recruitment on genetic patchiness in the urchinEchinometra mathaei in Western Australia

Enzyme polymorphisms in the sea urchinEchinometra mathaei were examined to test the relative influences of population turnover and patchiness in recruitment on genetic heterogeneity. We found that the total variance in allelic frequency among three populations separated by approximately 4 km at Rottnest Island, Western Australia (collected in February 1985) is as large as that among five additional samples collected over a distance of 1 300 km along the Western Australian coast in August 1987. This suggests that the forces causing genetic differentiation act on a local scale and occur in a single generation. A comparison of sites with different histories of recruitment indicates that the observed genetic differences among age groups are the result of prerecruitment effects, and that differences among sites reflect their individual histories of recruitment.     

Nestmate recognition and the genetic relatedness of nests in the ant Formica pratensis

Genetic relatedness of the mound-building ant Formica pratensis was determined by means of microsatellite DNA polymorphism, and its impact on nestmate recognition was tested in a population in Southern Sweden (Oeland). Recognition between nests was measured by testing aggression levels between single pairs of workers. The genetic distances of nests (Nei's genetic distance) and the spatial distance of nests were correlated and both showed a strong relation to the aggression behavior. Multiple regression analysis revealed a stronger impact of genetic relatedness rather than spatial distances on aggression behavior. Neighbouring nests were more closely related than distant nests, which may reflect budding as a possible spreading mechanism. The genetic distance data showed that nestmate recognition was strongly genetically influenced in F. pratensis. Received: 2 October 1997 / Accepted after revision: 10 January 1998     

High levels of genetic subdivision of marine and estuarine populations of the estuarine catfishCnidoglanis macrocephalus (Plotosidae) in southwestern Australia

The cobblerCnidoglanis macrocephalus (Valenciennes) is an endemic marine and estuarine catfish from southern Australia. Conflicting views on the degree of isolation of the estuarine populations underscore general questions about genetic divergence in coastal species. Although estuaries are widely recognized as ecologically important, little work has been done on their role in favouring genetic divergence. In order to estimate the extent of genetic subdivision among nearshore marine and estuarine populations, electrophoretic variation of enzymes was examined in seven marine and six estuarine populations of cobbler from sites spanning 1500 km along the southwest Australian coastline. Among all populations, the mean standardized variance in allelic frequencies (F _ST) for six polymorphic loci was 0.277, a high value comparable to those of other shallow-water teleosts whose life-history characteristics and habitat preferences restrict their dispersal capability. The pattern of genetic identities between populations showed divergence between west and south coast sites. Within these regional groups, however, there was substantial heterogeneity, much of which was associated with estuaries. Among all six estuarine sites, the averageF _ST was 0.333, 40% higher than the value of 0.237 for the marine sites. Low estimates of the genetically effective number of migrants suggest population subdivision between marine and estuarine environments and between similar habitat types. This study indicates the importance of habitat in affecting the connectedness of populations, even in apparently open marine systems.     

Genetic relatedness and eusociality: parasite-mediated selection on the genetic composition of groups

Summary Contrary to the expectations of kin selection theory, intracolony relatedness in eusocial insects is often low. We examined the idea that associations of low relatedness (high genetic variability) may be advantageous because of negative frequency-dependent selection on common host phenotypes by rapidly evolving parasites and pathogens. Using the natural host-parasite system of the bumble bee Bombus terrestris and its intestinal trypanosome Crithidia bombi, we investigated the transmission properties of parasites in host groups. Within naturally infested nests and in artificially constructed groups of workers, prevalence of infestation increased with time of exposure (Table 1). The susceptibility of isolated groups of workers to the parasites to which they were exposed differed with identity and natural infestation of their nest of origin (Table 2). In addition, those workers that were related to the individual introducing an infection to their group were more likely to become infested than were unrelated workers (Table 3). Although the bumble bee workers in experimental boxes appeared to differ in behavior toward kin and non-kin, making more physical contacts with kin, we found no discernible relationship between number of physical contacts and prevalence of infestation in a group. Therefore, we conclude that differences in parasite transmission reflected interactions among different host and parasite phenotypes. This system thus demonstrates the factors necessary for negative frequency-dependent selection by parasites on common host phenotypes - variability for susceptibility and infectiousness in host and parasite populations and similarity for these traits among related individuals. If, as we show here, high genetic relatedness within groups enhances parasite transmission, kin directed altruism may increase the risk of contracting parasites and infectious diseases. Therefore, parasites and pathogens may be an important force moderating the genetic structure of social groups. Offprint requests to: J.A. Shykoff at the present address     

Habitat patchiness and predation modify the distribution of a coral-dwelling damselfish

Fish abundance is often better predicted by microhabitat variables on continuous reefs than on isolated patch reefs. Although this was suggested to stem from reduced post-recruitment relocation, this has not been shown experimentally. We found the relationship between the presence of a coral-dwelling fish, Dascyllus marginatus, and the size of its coral host to differ between corals on continuous reefs and the sparsely distributed corals on sandy bottoms. Empty transplanted corals were colonized exclusively by new recruits when on the sandy bottom, and both by new recruits and post-recruitment dispersal of adults when on the continuous reef. New recruits settled predominantly into small corals, although analyses of recruitment patterns were confounded by low recruitment in the studied years. Both tank experiments and field survey data suggest that the presence of recruits in small corals is at least partially driven by predation by the dottyback, Pseudochromis olivaceus, which lives predominantly in large corals within both habitats. Consequently, we suggest that the relationship between fish presence and coral size differs between the habitats due to coral size dependent predation on recruits and variability in the importance of direct recruitment to replenish fish populations.     

High genetic relatedness among nestmate queens in the polygynous ponerine ant Gnamptogenys striatula in Brazil

Gnamptogenys striatula is a polygynous ant species, in which all workers are potentially able to mate. The reproductive status, relatedness and pedigree relationships among nestmate queens and winged females in a Brazilian population were investigated. We collected all the sexual females of 12 colonies (2–44 queens per colony, plus 2–18 winged females in 3 colonies). Dissections revealed that 98% of the queens were inseminated and that the queens in the most polygynous colonies did not lay equal numbers of eggs. The sexual females and a sample of the population were genotyped using eight microsatellite markers. Relatedness among nestmate queens was among the highest recorded to date (0.65±0.25), and tests of pedigree relationship showed that they were likely to be full-sisters, and sometimes cousins. Mated winged females were always full-sisters, the estimated genetically effective queen numbers were low and tests of pedigree relationship showed that only a few queens in the colony could be the mothers. These results suggest that the high queen-queen relatedness in polygynous colonies of G. striatula is maintained by an unusual mechanism: winged females are mostly produced by only one or a few queens, and these groups of full-sisters are recruited back into their original nest after mating. Received: 26 November 1999 / Revised: 7 September 2000 / Accepted: 7 September 2000     

Sibling recognition in thirteen-lined ground squirrels: effects of genetic relatedness,rearing association,and olfaction

Summary I investigated sibling-sibling recognition in captive thirteen-lined ground squirrels (Spermophilus tridecemlineatus) by cross-fostering lab-born pups shortly after birth. When young reached about 45 days of age, I observed dyadic interactions in a test arena of pairs from four relatedness X rearing groups, and recorded the frequency of exploratory encounters between individuals. Sibs-reared together and nonsibs-reared together exhibited significantly fewer exploratory encounters than either sibs-reared apart or nonsibsreared apart. Young reared together were equally exploratory, regadless of relatedness; similarly, young reared apart, whether they were sibs or nonsibs, showed similar levels of exploration. Thus, the differential treatment of siblings in the lab appears to be based on rearing association and not genetic relatedness per se. I interpret this recognition based on association (rearing familiarity) in the context of the species' social organization and compare my results on S. tridecemlineatus with similar studies on S. beldingi, S. parryii, and S. richardsonii. I also used an olfactory impairment technique (zinc sulfate) and found that differential treatment in thirteen-lined ground squirrels was influenced by olfactory cues.     

Environmental grain and polychaete species' diversity in a bathyal benthic community

Patterns of polychaete species' dispersion in the San Diego Trough, Southern California Continental Borderland, North Pacific Ocean, at a depth of 1230 m are analyzed. Samples consist of 4 replicate 0.25-m² box cores, each partitioned into twenty-five 0.01-m² subcores, and of 1 unpartitioned core. The sampler is biased but, when this bias is taken into account, few species show strong aggregation either between or within cores. If all species are considered together as replicates, intraspecific aggregation is detected between cores, but uniform dispersion dominates within cores. Patchiness or habitat partitioning on the assayed scales can thus account for little of the extremely high species' diversity observed. Apparently due to an environmental structure having a “grain” smaller than 0.01-m², the diversity of species assumed to be sedentary is higher than the diversity of species assumed to be mobile. One evidence of such structure is a negative covariance between paraonid abundance and density of “mudballs” constructed by a cirratulid. Consideration of these results suggests that either grain specialization or simultaneous, biogenic disequilibrium or both act on spatial scales smaller than 0.01-m² to maintain high polychaete species' diversity in the San Diego Trough.     

Phytoplankton patchiness and their role in the modelled productivity of a large, seasonally stratified lake

Phytoplankton concentration in Lake Kinneret (Israel) has varied up to 10-fold in space and time, with horizontal patches ranging from a couple of kilometres to a basin scale. Previous studies have used a 1D model to reproduce the temporal evolution of physical and biogeochemical variables in this lake. The question that arises then is how appropriate is a 1D approach to represent the dynamic of a spatially heterogeneous system, where there are non-linear dependencies between variables. Field data, a N-P-Z model coupled to both a 1D and a 3D hydrodynamic model, a 1D diffusion-reaction equation and scaling analysis are used to understand the role of spatial variability, expressed as phytoplankton patchiness, in the modelling of primary production. The analysis and results are used to investigate the effect of horizontal variability in the forcing and in the free mechanisms that affect the growth of patterns. The study shows that the use of averaged properties in a 1D approach may produce misleading results in the presence of localised patches, in terms of both concentration and composition of phytoplankton. The reason lies in the fact that the calibration process of ecological parameters in the 1D model appears to be site and process specific. That is, it depends on the pattern's characteristics and the underlying physical processes causing them. And this is a critical point for the success of numerical simulations under spatial variability. In this study, it is also shown that a length scale based on diffusion and growth rate of phytoplankton could be used as a criterion to assess the appropriateness of the 1D assumption.     

Disturbance and landscape dynamics in a changing world

Disturbance regimes are changing rapidly, and the consequences of such changes for ecosystems and linked social-ecological systems will be profound. This paper synthesizes current understanding of disturbance with an emphasis on fundamental contributions to contemporary landscape and ecosystem ecology, then identifies future research priorities. Studies of disturbance led to insights about heterogeneity, scale, and thresholds in space and time and catalyzed new paradigms in ecology. Because they create vegetation patterns, disturbances also establish spatial patterns of many ecosystem processes on the landscape. Drivers of global change will produce new spatial patterns, altered disturbance regimes, novel trajectories of change, and surprises. Future disturbances will continue to provide valuable opportunities for studying pattern-process interactions. Changing disturbance regimes will produce acute changes in ecosystems and ecosystem services over the short (years to decades) and long-term (centuries and beyond). Future research should address questions related to (1) disturbances as catalysts of rapid ecological change, (2) interactions among disturbances, (3) relationships between disturbance and society, especially the intersection of land use and disturbance, and (4) feedbacks from disturbance to other global drivers. Ecologists should make a renewed and concerted effort to understand and anticipate the causes and consequences of changing disturbance regimes.     

Modeling and data assessment of longitudinal salinity in a low-gradient estuarine river

Continuous data of vertical-profile salinity were analyzed for four stations located successively upriver in a macrotidal estuary, the lower St. Johns River (Northeast Florida, USA). The data analysis confirmed well-mixed salinity conditions in the river with at most 1.3 ppt of vertical variability at Dames Point (river km 20), where the main variations of salinity are along the longitudinal axis of the river. Given the well-mixed salinity conditions and dominant horizontal structure of salinity variations in the river, we present and apply a barotropic, two-dimensional modeling approach for hydrodynamic-salinity transport simulation in the lower St. Johns River. When properly forced by offshore surge, high-resolution wind fields and freshwater river inflows, the model replicated the salinity measurements remarkably well, including the separation into tidal and sub-tidal components. The data and model results show that, at times, offshore winds and surge can be more influential on longitudinal salinity variations than local winds over the river. We demonstrate the importance of using proper boundary conditions to force the model relative to the minimal sensitivity of the model to parameter adjustment of horizontal mixing and uncertainty-based perturbation of wind and inflow forcings.     

Diel flux of dissolved carbohydrate in a salt marsh and a simulated estuarine ecosystem

The concentrations of total dissolved carbohydrate (TCHO), monosaccharide (MCHO) and polysaccharide (PCHO) were followed over a total of ten diel cycles in a salt marsh and a 13 m³ seawater tank simulating an estuarine ecosystem. Their patterns are compared to those for total dissolved organic carbon (DOC), CO₂, pH, O₂, chlorophyll a, phaeopigments and solar radiation. During 5 of the 6 marsh studies, PCHO underwent periods of sustained accumulation starting in the late morning or early afternoon and continuing into the early evening. These periods possibly represent release of recently synthesized PCHO from phototrophs. similar patterns were not found in the tank although direct associations between TCHO and phaeopigment dynamics suggest that zooplankton excretion was an important source of dissolved carbohydrate. The numbers of planktonic bacteria determined in one tank study increased rapidly during a late morning PCHO pulse and varied inversely with PCHO throughout the afternoon and evening, indicating that they were able to respond rapidly and control natural substrate concentrations on a time scale of a few hours. MCHO fluctuated to a much lesser extent than PCHO at both locations with levels possibly maintained near the bacterial uptake threshold or in a closely regulated steady state. TCHO concentration changes over 2-to 3-h sampling intervals suggest very rapid net system release and uptake with summer rates frequently exceeding 30 g C l^-1h^-1 in the marsh and 20 g C l^-1 h^-1 in the tank.