Characterisation of microsatellite loci in the habitat-forming kelp, Ecklonia radiata (Phaeophyceae, Laminariales)

Habitat forming algae play an important role in the ecology of temperate reefs worldwide. Despite this, our understanding of levels of gene flow within and among populations of algae is largely limited to studies on intertidal species; we know comparatively little about important habitat-forming subtidal algae. Here, we develop eight polymorphic microsatellite markers for the characterisation of population genetic diversity and structure in the subtidal kelp, Ecklonia radiata. This large macroalga is the most abundant habitat-forming kelp on the subtidal rocky reefs of temperate Australia and New Zealand where it forms extensive forests that support an astounding diversity of associated taxa.     

Evolutionary Consequences of Dispersal Ability in Cactus-feeding Insects

Although gene flow is an important determinant of evolutionary change, the role of ecological factors such as specialization in determining migration and gene flow has rarely been explored empirically. To examine the consequences of dispersal ability and habitat patchiness on gene flow, migration rates were compared in three cactophagous longhorn beetles using coalescent analyses of mtDNA sequences. Analyses of covariance were used to identify the roles of dispersal ability and habitat distribution in determining migration patterns. Dispersal ability was a highly significant predictor of gene flow (p< 0.001), and was more important than any other factor. These findings predict that dispersal ability may be an import factor shaping both microevolutionary and macroevolutionary patterns; this prediction is borne out by comparisons of species diversity in cactus-feeding groups.     

Measures of gene flow in the Columbian ground squirrel

From analyses of published data and a review of the literature, I studied indirect and direct measures of gene flow among populations of Columbian ground squirrels, Spermophilus columbianus. New analyses were used to examine an allozyme data set (seven polymorphic loci) that had been collected by Zammuto and Millar (1985a) from six populations of ground squirrels that were spread over 183 km. G-tests indicated significant variation in allele frequencies among populations, but F-statistics revealed relatively little population differentiation (average F _ST=0.026). F _ST values were used to estimate rates of gene flow indirectly and indicated fairly high rates of gene flow (average N _e m=13.5). Recorded dispersal distances of individual ground squirrels were fairly short (most<4 km, maximum recorded distance was 8.5 km), and the minimum distance between populations used to create the allozyme data set was about 25 km. Thus, direct dispersal among the populations in the allozyme data set was highly unlikely. Small genetically effective populations may have experienced high rates of migration over short distances (about 43% of adults in local populations were immigrants), however, resulting in homogeneous allele frequencies over the geographic range. This explanation provides an alternative to invoking gene flow in the recent past to explain discrepancies between dispersal distances in the field and homogenization of allele frequencies over large ranges, Mammalian species that have virtually complete dispersal of subadult males from the natal area might be expected to exhibit relatively high rates of gene flow, regardless of actual dispersal distances. Genetically effective populations may be much smaller than more extensive ecological populations and experience higher rates of gene flow.     

Gene flow in a direct-developing,leaf litter frog between isolated mountains in the Taita Hills,Kenya

Amphibians are in decline worldwide, and high altitude tropical areas appear to be the worst affected. This is in stark contrast with current information we have on gene flow in amphibian populations which focus on temperate pond breeding species. Using AFLP markers, we show that a small, direct-developing, leaf litter frog from the Taita Hills in south–west Kenya (Schoutedenella xenodactyloides) has extended populations covering large areas (>3.5 km) of fragmented, forest habitat, uncharacteristic of typical amphibian models. Further, we demonstrate high levels of gene flow (F _ST < 0.065) through unsuitable dry savannah habitat which might otherwise be considered a barrier to dispersal. Landscape genetic analysis demonstrates a strong link between hydrologic features, and further highlights links between sites through specific catchments. We propose a model of passive-active dispersal for the Dwarf Squeaker, S. xenodactyloides, which features passive downhill and active uphill movements over large areas, contrasting with limited cross slope movements. Our study highlights the importance of the diverse reproductive strategies of the Amphibia when considering dispersal and gene flow, and hence conservation management.     

The role of spatial and ontogenetic morphological variation in the expansion of the geographic range of the tropical brown alga, Turbinaria ornata

Like many reefs worldwide, reefs in French Polynesia are experiencing a shift from coral-dominated to algal-dominated systems. The macroalga Turbinaria ornata comprises the majority of the increasing algal biomass on the barrier reefs surrounding these islands, and its distribution is increasing throughout this region. Aspects of the ecomorphology of Turbinaria make it ideally suited to thrive under the physical conditions found across barrier reefs throughout French Polynesia. Spatial morphological variation allows Turbinaria to produce morphotypes that are suited either to the calm, unidirectional, slowly flowing water in the backreef or to the high-energy wave-driven flow of the forereef. Backreef plants are flexible and produce airbladders that make them buoyant, whereas forereef plants are not buoyant, but strong and stiff. Production of bladders and resulting buoyancy has been found to be a phenotypically plastic trait in response to movement of water and confers advantages to backreef plants and plays an important role in dispersal. Ontogenetic variation of buoyancy, material properties, and reproductive capacity is part of a dispersal strategy whereby fertile, buoyant fronds drift between oceanic islands and form new populations, thereby contributing to the recent expansion of range of T. ornata across French Polynesia.     

Spatial genetic structure and restricted gene flow in a functionally dioecious fig, <Emphasis Type="Italic">Ficus pumila</Emphasis> L. var. <Emphasis Type="Italic">pumila</Emphasis> (Moraceae)

The mutualism between fig plants and fig wasps has been recognized as one of the most specialized systems of symbiosis. Figs are pollinated by their highly specific pollinating fig wasps, and the pollinating fig wasps are raised within the syconia of figs. Recent studies indicated a difference between monoecious and dioecious figs in the dispersal range of pollinating wasps, which has potential consequences for gene flow. In this study, we detected the gene-flow pattern of the dioecious climbing fig, Ficus pumila L. var. pumila, at both local and regional scales. At the local scale, spatial autocorrelation analysis indicated strong genetic structure at short distances, a pattern of limited gene flow. This result was also supported by a high inbreeding coefficient (F _IS = 0.287) and significant substructuring (F _ST = 0.060; P < 0.001). Further analysis indicated that the effective gene dispersal range was 1,211 m, and the relative contribution of seed dispersal was smaller than that of pollen dispersal. The inferred effective range of pollen dispersal ranged from 989 to 1,712 m, while the effective seed dispersal range was less than 989 m. Lack of long-distance dispersal agents may explain the limited seed dispersal. The high density of receptive fig trees was the most likely explanation for limited pollen dispersal, and the position of syconia and relatively low wind speed beneath the canopy may contribute to this phenomenon. At the regional scale, significant negative correlations (kinship coefficient F _ij ranging from −0.038 to −0.071) existed in all comparisons between the studied population and other populations, and the assignment test grouped almost all individuals of the studied population into a distinct cluster. Asynchronous flowering on the regional scale, which provides a barrier for the pollinating wasps to fly from the studied population to the other populations, is probably responsible for the limited gene flow on the regional scale.     

Reduced performance of native infauna following recruitment to a habitat-forming invasive marine alga

Despite well-documented negative impacts of invasive species on native biota, evidence for the facilitation of native organisms, particularly by habitat-forming invasive species, is increasing. However, most of these studies are conducted at the population or community level, and we know little about the individual fitness consequences of recruitment to habitat-forming invasive species and, consequently, whether recruitment to these habitats is adaptive. We determined the consequences of recruitment to the invasive green alga Caulerpa taxifolia on the native soft-sediment bivalve Anadara trapezia and nearby unvegetated sediment. Initially, we documented the growth and survivorship of A. trapezia following a natural recruitment event, to which recruitment to C. taxifolia was very high. After 12 months, few clams remained in either habitat, and those that remained showed little growth. Experimental manipulations of recruits demonstrated that all performance measures (survivorship, growth and condition) were significantly reduced in C. taxifolia sediments compared to unvegetated sediments. Exploration of potential mechanisms responsible for the reduced performance in C. taxifolia sediments showed that water flow and water column dissolved oxygen (DO) were significantly reduced under the canopy of C. taxifolia and that sediment anoxia was significantly higher and sediment sulphides greater in C. taxifolia sediments. However, phytoplankton abundance (an indicator of food supply) was significantly higher in C. taxifolia sediments than in unvegetated ones. Our results demonstrate that recruitment of native species to habitat-forming invasive species can reduce growth, condition and survivorship and that studies conducted at the community level may lead to erroneous conclusions about the impacts of invaders and should include studies on life-history traits, particularly juveniles.     

Dispersal depends on body condition and predation risk in the semi‐aquatic insect,Notonecta undulata

Dispersal is the movement of organisms across space, which has important implications for ecological and evolutionary processes, including community composition and gene flow. Previous studies have demonstrated that dispersal is influenced by body condition; however, few studies have been able to separate the effects of body condition from correlated variables such as body size. Moreover, the results of these studies have been inconsistent with respect to the direction of the relationship between condition and dispersal. We examined whether body condition influences dispersal in backswimmers (Notonecta undulata). We also tested whether an interaction between body condition and predation risk (another proximate factor that influences dispersal) could contribute to the previously observed inconsistent relationship between condition and dispersal. We imposed diet treatments on backswimmers in the laboratory, and measured the effects of food availability on body condition and dispersal in the field. We found that dispersal was a positive function of body condition, which may have important consequences for population characteristics such as the rate of gene flow and population growth. However, the effects of body condition and predation risk were additive, not interactive, and therefore, our data do not support the hypothesis that the interaction between condition and predation risk contributes to the inconsistency in the results of previous condition‐dependent dispersal studies.     

High gene flow between populations of Macrotermes michaelseni (Isoptera, Termitidae)

Termite alates are thought to be poor active flyers, and this should lead to considerable genetic differentiation on small spatial scales. However, using four microsatellite loci for the termite Macrotermes michaelseni we found low values of genetic differentiation (F_ST) across a spatial scale of even more than 50 km. Genetic differentiation between populations increased with spatial distance up to 50 km. Furthermore, up to this distance, the scatter around the linear regression of genetic differentiation versus spatial distance increased with spatial distance. This suggests that across such spatial distances gene flow and genetic drift are of about equal importance, and near equilibrium. Using a regional F_ST as well as the distance between populations with non-significant F_ST-values (up to 25 km), gene flow is sufficiently high so that populations may be regarded as panmictic on spatial scales of 25 to 50 km. The apparent contradiction between dispersal distances observed in the field and estimates of gene flow from genetic markers may be due to the masses of swarming alates. Assuming a leptokurtic distribution of dispersal distances, atleast some alates are expected to travel considerable distances, most likely by passive drift. Received 25 January 2005; revised 11 April 2005; accepted 26 April 2005.     

Maintenance of high pollen dispersal in Eucalyptus wandoo, a dominant tree of the fragmented agricultural region in Western Australia

Pollen dispersal was investigated in five remnant populations of Eucalyptus wandoo, a dominant insect-pollinated tree in the fragmented agricultural region of southern Western Australia. Paternity analysis using six microsatellite loci identified a pollen source for 45% of seedlings, and the remainder were assumed to have arisen from pollen sources outside the stands. Outcrossing was variable, ranging from 52 to 89%, and long distance pollen dispersal was observed in all populations with up to 65% of pollen sourced from outside the populations over distances of at least 1 km. Modelling dispersal functions for pollination events within the two larger populations showed little difference between the four two-parameter models tested and indicated a fat-tailed dispersal curve. Similarity of direct and indirect historical estimates of gene flow indicates maintenance of gene flow at levels experienced prior to fragmentation. The study revealed extensive long distance pollen dispersal in remnant patches of trees within a fragmented agricultural landscape in the southern temperate region and highlighted the role of remnant patches in maintaining genetic connectivity at the landscape scale.     

Small fish,large river: Surprisingly minimal genetic structure in a dispersal‐limited,habitat specialist fish

Genetic connectivity is expected to be lower in species with limited dispersal ability and a high degree of habitat specialization (intrinsic factors). Also, gene flow is predicted to be limited by habitat conditions such as physical barriers and geographic distance (extrinsic factors). We investigated the effects of distance, intervening pools, and rapids on gene flow in a species, the Tuxedo Darter (Etheostoma lemniscatum), a habitat specialist that is presumed to be dispersal‐limited. We predicted that the interplay between these intrinsic and extrinsic factors would limit dispersal and lead to genetic structure even at the small spatial scale of the species range (a 38.6 km river reach). The simple linear distribution of E. lemniscatum allowed for an ideal test of how these factors acted on gene flow and allowed us to test expectations (e.g., isolation‐by‐distance) of linearly distributed species. Using 20 microsatellites from 163 individuals collected from 18 habitat patches, we observed low levels of genetic structure that were related to geographic distance and rapids, though these factors were not barriers to gene flow. Pools separating habitat patches did not contribute to any observed genetic structure. Overall, E. lemniscatum maintains gene flow across its range and is comprised of a single population. Due to the linear distribution of the species, a stepping‐stone model of dispersal best explains the maintenance of gene flow across its small range. In general, our observation of higher‐than‐expected connectivity likely stems from an adaptation to disperse due to temporally unstable and patchy habitat.     

Turbinaria ornata (Phaeophyceae) varies size and strength to maintain environmental safety factor across flow regimes

Water motion in coastal areas can produce hydrodynamic forces that damage or dislodge benthic macroalgae if the tissues of macroalgae are not sufficiently strong. Some macroalgae vary their morphology and strength in response to ambient water motion, but little is known of how morphology and strength of macroalgae change relative to one another across flow regimes. Here, we use Turbinaria ornata, an ecologically important macroalga, to study how both the morphology and strength of macroalgae vary with ambient water motion. Typically, T. ornata exhibits weakening of its stipe when sexually mature, leading to breakage from the substratum and dispersal, which is beneficial for reproduction. Across three flow regimes, adult T. ornata increased its size but decreased its strength as water motion decreased. However, the strength of T. ornata relative to the maximum hydrodynamic forces it is expected to encounter (the environmental safety factor) did not differ between flow regimes. Our results showed that T. ornata can conform to its local flow habitat by varying both size and strength, similar to other macroalgae. Varying multiple traits between flow regimes suggested that T. ornata is capable of surviving a wide range of flow conditions, which may permit more control over the timing of its weakening, breakage from the substratum, and dispersal, even with future increases in flow velocities (e.g., large waves from storms) that are expected to occur frequently with climate change.     

Gene Flow,Risk Assessment and the Environmental Release of Transgenic Plants

The release of genetically modified plants is governed by regulations that aim to provide an assessment of potential impact on the environment. One of the most important components of this risk assessment is an evaluation of the probability of gene flow. In this review, we provide an overview of the current literature on gene flow from transgenic plants, providing a framework of issues for those considering the release of a transgenic plant into the environment. For some plants gene flow from transgenic crops is well documented, and this information is discussed in detail in this review. Mechanisms of gene flow vary from plant species to plant species and range from the possibility of asexual propagation, short- or long-distance pollen dispersal mediated by insects or wind and seed dispersal. Volunteer populations of transgenic plants may occur where seed is inadvertently spread during harvest or commercial distribution. If there are wild populations related to the transgenic crop then hybridization and eventually introgression in the wild may occur, as it has for herbicide resistant transgenic oilseed rape (Brassica napus). Tools to measure the amount of gene flow, experimental data measuring the distance of pollen dispersal, and experiments measuring hybridization and seed survivability are discussed in this review. The various methods that have been proposed to prevent gene flow from genetically modified plants are also described. The current “transgenic traits” in the major crops confer resistance to herbicides and certain insects. Such traits could confer a selective advantage (an increase in fitness) in wild plant populations in some circumstances, were gene flow to occur. However, there is ample evidence that gene flow from crops to related wild species occurred before the development of transgenic crops and this should be taken into account in the risk assessment process.     

Combining Genetic and Demographic Data for the Conservation of a Mediterranean Marine Habitat-Forming Species

The integration of ecological and evolutionary data is highly valuable for conservation planning. However, it has been rarely used in the marine realm, where the adequate design of marine protected areas (MPAs) is urgently needed. Here, we examined the interacting processes underlying the patterns of genetic structure and demographic strucuture of a highly vulnerable Mediterranean habitat-forming species (i.e. Paramuricea clavata (Risso, 1826)), with particular emphasis on the processes of contemporary dispersal, genetic drift, and colonization of a new population. Isolation by distance and genetic discontinuities were found, and three genetic clusters were detected; each submitted to variations in the relative impact of drift and gene flow. No founder effect was found in the new population. The interplay of ecology and evolution revealed that drift is strongly impacting the smallest, most isolated populations, where partial mortality of individuals was highest. Moreover, the eco-evolutionary analyses entailed important conservation implications for P. clavata. Our study supports the inclusion of habitat-forming organisms in the design of MPAs and highlights the need to account for genetic drift in the development of MPAs. Moreover, it reinforces the importance of integrating genetic and demographic data in marine conservation.     

ABSENCE OF A LARGE BROWN MACROALGA ON URBANIZED ROCKY REEFS AROUND SYDNEY,AUSTRALIA, AND EVIDENCE FOR HISTORICAL DECLINE1

Loss of habitat‐forming algae is increasingly prevalent in temperate marine ecosystems. Here, we document absence of an important habitat‐forming macroalga, Phyllospora comosa (Labill.) C. Agardh, along an urbanized coast in New South Wales (NSW), Australia. Dense Phyllospora canopies were common on shallow sublittoral reefs north and south of Sydney. In contrast, we did not find a single individual along ～70 km of rocky coastline in the Sydney metropolitan region, despite historical evidence to suggest that it was very common half a century ago. Recolonization of this important habitat‐forming alga has not occurred on Sydney reefs despite improved water quality, protection of its habitat, and frequent long‐distance dispersal of Phyllospora wrack. While there are obvious limitations, historical information can be useful for identifying potential shifts in community structure to increase our understanding of contemporary ecological patterns.     

Landscape genetics of a top neotropical predator

Habitat loss and fragmentation as a consequence of human activities is a worldwide phenomenon and one of the major threats to global biodiversity. Habitat loss and fragmentation is particularly a concern in the biodiverse tropics, where deforestation is occurring at unprecedented rates. Although insects are one of the most diverse and functionally important groups in tropical ecosystems, the quantitative effect of landscape features on their gene flow remains unknown. Here, we used a robust landscape genetics approach to quantify the effect of ten landscape features (deforestation, mature forests, other forest types, the River Chagres, streams, stream banks, roads, sea, lakes and swamps) and interactions between them, on the gene flow of a neotropical forest keystone species, the army ant Eciton burchellii. The influence of landscape on E. burchellii's gene flow reflected the different dispersal capability of its sexes; aerial for males and pedestrian for females, and the different depths of population history inferred from microsatellites and mitochondrial DNA. In contrast to the gene flow‐facilitating effect of mature forests, deforested areas were found to be strong barriers for E. burchellii's gene flow. Other forest types were found to be gene flow facilitators but only when interacting with mature secondary forests, therefore indicating the importance of mature forests for the survival of E. burchelii and its associate species. The River Chagres was identified as a major historical gene flow barrier for E. burchellii, suggesting that an important loss of connectivity may occur because of large artificial waterways such as the Panama Canal.     

Evidence of partial anadromy and resident-form dispersal bias on a fine scale in populations of Oncorhynchus mykiss

We examine sympatric anadromous (steelhead) and nonanadromous (resident) rainbow trout (Oncorhynchus mykiss) from neighboring locations to test three hypotheses: (1) the sympatric life history types are not genetically different; (2) fine-scale dispersal is the same for both sexes, and (3) fine-scale dispersal is the same for steelhead and resident individuals. Data from 13 microsatellite loci reveal no genetic difference between sympatric steelhead and resident O. mykiss but moderate population structure (F _ST=0.019–0.028) between adjacent samples, regardless of life history type. Our results provide further evidence of partial anadromy and suggest that geographic proximity and genetic history, more than migratory type, should be considered when identifying populations for use in restoration of local genetic diversity. We find evidence of resident-form dispersal bias on a fine spatial scale, however, we find no evidence that fine-scale dispersal varies by gender. Conservation strategies should aim to maintain resident and anadromous forms when they occur in sympatry, as they may be important in facilitating gene flow on small and large spatial scales, respectively.     

Dispersal Behavior, Boldness, and the Link to Invasiveness: A Comparison of Four Gambusia Species

Dispersal is a key element of a species' invasiveness. Although considerable work has addressed how dispersal influences the pattern of spatial spread of invading organisms, few studies investigate whether invasive species are in fact better dispersers than either the species they displace or less successful invaders. Recent work suggests that variation in dispersal may be due to variation in an underlying behavioral trait, boldness. Our study examined the link between dispersal, boldness, and invasiveness by comparing the dispersal characteristics and refuge use of two invasive Gambusia species to two congeners in experimental streams. The streams consisted of a series of pools (no flow) connected to a flowing channel. For each species, small groups of females were released at the middle pool, and their movement and activity were recorded over a 1-h period. We found invasive Gambusia to be more likely to disperse out of the introductory pool, to disperse sooner, to travel a greater distance in the artificial streams, and thus to exhibit greater dispersal tendencies than their close relatives. Among the invasives, Gambusia affinis had a greater dispersal tendency than G. holbrooki. We suspect this result indicates variation in the contribution of dispersal to the relative invasiveness of these species. Certain dispersal measures were correlated to time spent out of refuge, although invasive Gambusia and their relatives did not differ in the predicted manner. These results argue for the greater incorporation of experimental approaches and analyses of behavioral mechanisms in the study of invasive species.     

Genetic variability in two hydrophilous species ofPotamogeton,P. pectinatus andP. filiformis (Potamogetonaceae)

Genetic variability inPotamogeton pectinatus andP. filiformis was studied by means of isozymes. The overall levels of variability were similar to some other well studied hydrophilous species, and were shown to be distributed more between than within populations. This partitioning of variability was attributed to three main factors. (1) Clonal growth (as measured by the frequency of multi-enzyme phenotypes) was shown to be a major factor in both species, although more important inP. pectinatus. (2) Low levels of sexual reproduction were shown to be a likely contributor to the partitioning of variability inP. pectinatus; such reproduction is probably limited by seedling recruitment rather than by infrequent flowering. (3) Geographical isolation was indicated as a factor regulating gene flow at distances of more than about 1000 km in both species, with the data suggesting that dispersal between populations is mainly by seed rather than by vegetative means.