Seasonal dimorphism in the horny bills of sparrows

Bill size is often viewed as a species‐specific adaptation for feeding, but it sometimes varies between sexes, suggesting that sexual selection or intersexual competition may also be important. Hypotheses to explain sexual dimorphism in avian bill size include divergence in feeding niche or thermoregulatory demands, intrasexual selection based on increased competition among males, or female preference. Birds also show seasonal changes in bill size due to shifts in the balance between growth rate and wear, which may be due to diet or endogenous rhythms in growth. Insight into the function of dimorphism can be gained using the novel approach of digital x‐ray imaging of museum skins to examine the degree to which the skeletal core or the rhamphotheca contribute to overall dimorphism. The rhamphotheca is ever‐growing and ever‐wearing, varying in size throughout life; whereas the skeletal core shows determinant growth. Because tidal marsh sparrows are more dimorphic in bill size than related taxa, we selected two marsh taxa to investigate dimorphism and seasonality in the size of the overall bill, the skeletal core, and the rhamphotheca. Bill size varied by sex and season, with males having larger bills than females, and bill size increasing from nonbreeding to breeding season more in males. Skeletal bill size varied with season, but not sex. The rhamphotheca varied primarily with sex; males had a larger rhamphotheca (corrected for skeletal bill size), which showed a greater seasonal increase than females. The rhamphotheca, rather than the skeletal bill, was responsible for sexual dimorphism in overall bill size, which was particularly well developed in the breeding season. The size of the rhamphotheca may be a condition‐based character that is shaped by sexual selection. These results are consistent with the evidence that bill size is influenced by sexual selection as well as trophic ecology.     

High summer temperature explains bill size variation in salt marsh sparrows

Physiological factors are rarely proposed to account for variation in the morphology of feeding structures. Recently, bird bills have been demonstrated to be important convective and radiant heat sinks. Larger bills have greater surface area than smaller bills and could serve as more effective thermoregulatory organs under hot conditions. The heat radiating function of bills should be more important in open habitats with little shade and stronger convective winds. Furthermore, as a means of dumping heat without increasing water loss through evaporation, bills might play a particularly important thermoregulatory role in heat loss in windy habitat where fresh water is limited. North American salt marshes provide a latitudinal gradient of relatively homogeneous habitat that is windy, open, and fresh‐water limited. To examine the potential role of thermoregulation in determining bill size variation among ten species or subspecies of tidal marsh sparrows, we plotted bill size against maximum summer and minimum winter temperatures. Bill surface areas increases with summer temperature, which explained 82–89% of the variance (depending upon sex) when we controlled for genus membership. Latitude alone predicted bill surface area much more poorly than summer temperature, and winter temperatures explained < 10% of the variance in winter bill size. Tidal marsh sparrow bill morphology may, to a large degree, reflect the role of the bill in expelling excess body heat in these unbuffered, fresh‐water‐limited environments. This new example of Allen's rule reaffirms the importance of physiological constraints on the evolution of vertebrate morphologies, even in bird bills, which have conventionally been considered as products of adaptation to foraging niche.     

Heat Loss May Explain Bill Size Differences between Birds Occupying Different Habitats

Background

Research on variation in bill morphology has focused on the role of diet. Bills have other functions, however, including a role in heat and water balance. The role of the bill in heat loss may be particularly important in birds where water is limiting. Song sparrows localized in coastal dunes and salt marsh edge (Melospiza melodia atlantica) are similar in size to, but have bills with a 17% greater surface area than, those that live in mesic habitats (M. m. melodia), a pattern shared with other coastal sparrows. We tested the hypotheses that sparrows can use their bills to dissipate “dry” heat, and that heat loss from the bill is higher in M. m. atlantica than M. m. melodia, which would indicate a role of heat loss and water conservation in selection for bill size.

Methodology/Principal Findings

Bill, tarsus, and body surface temperatures were measured using thermal imaging of sparrows exposed to temperatures from 15–37°C and combined with surface area and physical modeling to estimate the contribution of each body part to total heat loss. Song sparrow bills averaged 5–10°C hotter than ambient. The bill of M. m atlantica dissipated up to 33% more heat and 38% greater proportion of total heat than that of M. m. melodia. This could potentially reduce water loss requirements by approximately 7.7%.

Conclusions/Significance

This >30% higher heat loss in the bill of M. m. atlantica is independent of evaporative water loss and thus could play an important role in the water balance of sparrows occupying the hot and exposed dune/salt marsh environments during the summer. Heat loss capacity and water conservation could play an important role in the selection for bill size differences between bird populations and should be considered along with trophic adaptations when studying variation in bill size.     

Divergent selection and drift shape the genomes of two avian sister species spanning a saline–freshwater ecotone

The role of species divergence due to ecologically based divergent selection—or ecological speciation—in generating and maintaining biodiversity is a central question in evolutionary biology. Comparison of the genomes of phylogenetically related taxa spanning a selective habitat gradient enables discovery of divergent signatures of selection and thereby provides valuable insight into the role of divergent ecological selection in speciation. Tidal marsh ecosystems provide tractable opportunities for studying organisms' adaptations to selective pressures that underlie ecological divergence. Sharp environmental gradients across the saline–freshwater ecotone within tidal marshes present extreme adaptive challenges to terrestrial vertebrates. Here, we sequence 20 whole genomes of two avian sister species endemic to tidal marshes—the saltmarsh sparrow (Ammospiza caudacutus) and Nelson's sparrow (A. nelsoni)—to evaluate the influence of selective and demographic processes in shaping genome‐wide patterns of divergence. Genome‐wide divergence between these two recently diverged sister species was notably high (genome‐wide F_ST = 0.32). Against a background of high genome‐wide divergence, regions of elevated divergence were widespread throughout the genome, as opposed to focused within islands of differentiation. These patterns may be the result of genetic drift resulting from past tidal march colonization events in conjunction with divergent selection to different environments. We identified several candidate genes that exhibited elevated divergence between saltmarsh and Nelson's sparrows, including genes linked to osmotic regulation, circadian rhythm, and plumage melanism—all putative candidates linked to adaptation to tidal marsh environments. These findings provide new insights into the roles of divergent selection and genetic drift in generating and maintaining biodiversity.     

Population history and the selective landscape shape patterns of osmoregulatory trait divergence in tidal marsh Savannah sparrows (Passerculus sandwichensis)

A persistent challenge in making associations between phenotypic and environmental variation is understanding how ecological factors and demographic history interact to shape adaptive outcomes. Evaluating the degree to which conspecific populations exposed to similar environmental pressures respond in parallel provides a powerful framework for addressing this challenge. We took this comparative approach with multiple populations of Savannah sparrows (Passerculus sandwichensis) found in tidal marshes along the Pacific coast of North America. The high salinities characterizing tidal marshes select for increased osmoregulatory performance and salinity tolerance. We collected data on physiological traits associated with osmoregulatory performance from 10 tidal marsh and three freshwater-adapted interior populations to evaluate the degree of parallel divergence across populations. All traits showed differences in the magnitude of divergence, but only total evaporative water loss (TEWL) showed differences in the direction of divergence. The drivers of these differences in both the magnitude and direction of divergence varied among traits. For kidney morphology and TEWL, patterns of divergence were best explained by variation in immigration rate from interior populations. Maximum temperature was the best predictor of variation in urine excretion ability, and both gene flow and temperature contributed to variation in plasma osmolality. Finally, analysis of multitrait divergence patterns indicated that differences in the direction of divergence were best explained by population genetic structure, whereas differences in the magnitude of divergence were explained by environmental differences. Together these results show that the influences of demography and the selective landscape can manifest themselves differently across functionally integrated traits.     

Genotype‐environment associations support a mosaic hybrid zone between two tidal marsh birds

Local environmental features can shape hybrid zone dynamics when hybrids are bounded by ecotones or when patchily distributed habitat types lead to a corresponding mosaic of genotypes. We investigated the role of marsh‐level characteristics in shaping a hybrid zone between two recently diverged avian taxa – Saltmarsh (Ammodramus caudacutus) and Nelson's (A. nelsoni) sparrows. These species occupy different niches where allopatric, with caudacutus restricted to coastal marshes and nelsoni found in a broader array of wetland and grassland habitats and co‐occur in tidal marshes in sympatry. We determined the influence of habitat types on the distribution of pure and hybrid sparrows and assessed the degree of overlap in the ecological niche of each taxon. To do this, we sampled and genotyped 305 sparrows from 34 marshes across the hybrid zone and from adjacent regions. We used linear regression to test for associations between marsh characteristics and the distribution of pure and admixed sparrows. We found a positive correlation between genotype and environmental variables with a patchy distribution of genotypes and habitats across the hybrid zone. Ecological niche models suggest that the hybrid niche was more similar to that of A. nelsoni and habitat suitability was influenced strongly by distance from coastline. Our results support a mosaic model of hybrid zone maintenance, suggesting a role for local environmental features in shaping the distribution and frequency of pure species and hybrids across space.     

Tidal‐flow restoration provides little nesting habitat for a globally vulnerable saltmarsh bird

Tidal marshes are among the most threatened habitats on Earth because of their limited natural extent, a long history of human drainage and modification, and anticipated future sea‐level rise. Tidal marshes also provide services to humans and support species of high conservation interest. Consequently, millions of dollars have been spent on tidal marsh restoration throughout North America. Southern New England has a long history of tidal marsh restorations, often focused on removal of the invasive plant Phragmites australis. Working in 18 Connecticut marshes, we examined the bird community in 21 plots in restoration sites and 19 plots in reference sites. Restoration plots were divided into those in marshes where management involved restoring tidal flow and those where direct Phragmites control (e.g. cutting, herbicide) was used. Saltmarsh sparrows Ammodramus caudacutus, which are considered globally vulnerable to extinction, were less common where tidal flow had been restored than at reference sites and nested in only one of 14 tidal‐flow restoration plots. No abundance differences were found for large wading birds, willets Tringa semipalmata, or seaside sparrows Ammodramus maritimus. Vegetation at sites where tidal flow had been restored showed characteristics typical of lower‐elevation marsh, which is unsuitable for nesting saltmarsh sparrows. We conclude that, although tidal‐flow restorations in Connecticut control Phragmites and restore native saltmarsh vegetation, they produce conditions that are largely unsuitable for one of the highest conservation priority species found in eastern U.S. salt marshes.     

Hydraulic Geometry: A Geomorphic Design Tool for Tidal Marsh Channel Evolution in Wetland Restoration Projects

Empirical hydraulic geometry relationships for tidal marsh channels are a practical geomorphically based design tool that can assist in the planning of tidal wetland restoration projects. This study provides hydraulic geometry relationships for predicting the depth, width, and cross‐sectional area of mature tidal channels as functions of contributing marsh area or tidal prism. The relationships are based on data from San Francisco Bay coastal salt marshes ranging in size from 2 to 5,700 ha. These hydraulic geometry relationships refine and expand on earlier relationships. Relationships for mature marshes can be used to predict the direction and rate of evolution of a channel in an immature or perturbed marsh system. Channel evolution data for three youthful tidal channels, ages 4 to 13 years, suggest that the channels are converging on their predicted equilibrium morphology. Two channels are eroding in response to significant increases in upstream tidal prism. They have enlarged first by deepening, in one case after 13 years to beyond the predicted equilibrium depth, and then widening through slumping of the channel banks. The third channel, a new one forming in a depositional mudflat, is converging on its equilibrium morphology after 13 years but will likely take several decades to equilibrate.     

Morphological adaptation influences the evolution of a mating signal

Theory predicts that forces of natural selection can reduce the intensity of sexually selected traits. In this study, I investigate how morphological adaptation to feeding ecology influences a mating signal. In birds, changes in feeding ecology can cause rapid divergence in bill morphology. Because bills are also important for song production, feeding ecology may influence song divergence. During song, birds can rapidly change vocal tract resonance using bill movement, yet are constrained in rate and magnitude of bill movements resulting in a trade-off between trill rate and frequency bandwidth. Male swamp sparrows vary in their ability to produce rapid, broad-band trills and females prefer more physically demanding songs. Populations of swamp sparrows adapted to the feeding ecology of tidal marshes have larger bills than inland populations. Larger bills should increase the constraints of producing rapid, broad-band trills allowing for a test of how changes in feeding ecology affect a feature of song used in mate choice. I found significant differences in acoustic features of song consistent with the hypothesis that coastal males are less able to meet the physical demands of song production because of the constraints of having larger bills. As possible compensation for decreases in song performance, coastal populations exhibit an increase in song complexity. These changes support the current model of how motor constraints influence song production and suggest a mechanism by which feeding ecology can influence signal evolution.     

Supratidal foraminifera as ecological indicators in anthropically modified wetlands (Lagoon of Venice, Italy)

In transitional environments, the intertidal zones represent a peculiar case characterized by halophile vegetation and by a low diversity benthic community. On these areas just a few particular foraminiferal species, a class of Protoctista secreting a shell called test, can survive for a certain time out of water. They are distributed in well-defined vertical zonations with respect to mean sea level and they correspond to analogous marsh floral zonations. In particular, the Trochammina macrescens Brady + Trochammina inflata (Montagu) association characterizes the salt marsh zone above mean high water level. The potential of these taxa as bioindicators is tested, since their presence-absence-dominance differentiates the subtidal/supratidal environments. Over the last few centuries, various engineering works generated major physical changes in the Venetian Lagoon. These changes affected the natural evolution of the intertidal morphologies, the surface of which is decreasing. In an attempt to reverse this tendency, numerous artificial salt marshes have been constructed and more are under construction. In this study, the Mazzorbo artificial salt marsh, built during the second half of 1999, is considered. On its surface, 16 samples were collected along a transect line in May 2008 to verify the ecological role of this salting within the lagoon ecosystem. The sediment grain size distribution of the salt marsh reflects the dissipative role of the tide and the effect of sediment transport due to the wave and tidal action. However, the presence of only a few Trochammina individuals shows that the foraminiferal fauna did not recognise this morphology as a salt marsh. The lack of Trochammina colonisation can be related to the excessive elevation of the salt marsh surface. This hypothesis is confirmed by the lack of the salt-tolerant plant Spartina. The unsuccessful colonisation by the foraminifera seems to indicate that this artificial salting does not have the natural dynamism of the intertidal morphologies and it may only be classified as land recovery. The supratidal foraminiferal taxa can act as an ecological indicator: through their observation it is possible to verify whether an artificial salt marsh accomplishes its task of functioning as an ecological unit with the community of organisms.     

Abundance and diversity of tidal marsh plants along the salinity gradient of the San Francisco Estuary: implications for global change ecology

From 2003 through 2005, tidal marsh plant species diversity and abundance on historically surveyed vegetation transects along the salinity gradient of the San Francisco Estuary were investigated to establish empirical relationships between plant distributions and environmental conditions, and furthermore to examine and predict past and future plant distribution changes. This study suggests that for most species, salinity is the primary control on plant distribution. Thus, ongoing changes in estuarine conditions (increasing sea level and salinity) are resulting in a complex mix of plant distribution changes. On the low marsh, where sediment salinity is similar to that of ambient water, halophytic species are replacing salt-intolerant taxa. However, on marsh plains, where increased tidal flooding is moderating high salinity (concentrated by evaporation), halophytic “high marsh” species are being replaced by salt-intolerant “low marsh” taxa. Thus, future changes in plant distributions will hinge on whether marsh sediment accumulation keeps pace with sea level rise.     

Assortative mating and patterns of inheritance indicate that the three crossbill taxa in Scotland are species

There are three breeding taxa of crossbills in highland Scotland, the common crossbill Loxia curvirostra , parrot crossbill L. pytyopsittacus and endemic Scottish crossbill L. scotica . These taxa show no genetic differentiation in neutral DNA (microsatellites and mitochondrial DNA), but do show some differentiation in morphology and have distinct calls. Mating patterns and bill size inheritance were examined to test four different hypotheses for their genetic similarity: (1) the three taxa belong to the same species and differences in bill sizes are caused by phenotypic plasticity, (2) the taxa are a result of genetic polymorphism at a single locus, (3) the taxa largely act as species but occasionally cross-breed, such that gene flow results in genetic homogenisation of part of the genome, and (4) the taxa are fully reproductively isolated species but diverged only recently, such that insufficient time has passed for neutral DNA to differentiate by genetic drift. Hypotheses 1 and 2 could be excluded because wild crossbills mated assortatively with respect to bill depth (a measure of bill size) and bill depth was highly heritable; narrow-sense heritabilities of 0.58 for female and 0.71 for male Scottish crossbills. Only two of 46 pairs (4.3%) were not assortative by bill size. Pairing was strongly assortative by flight calls (50 out of 52 pairs, 96.2%) and excitement calls (88 out of 93 pairs, 94.6%). Therefore, the three crossbill taxa do not cross-mate freely, nor do they seem totally reproductively isolated (excluding hypothesis 4). Thus, the results mostly favour hypothesis 3. The small amount of gene flow among crossbill taxa may contribute to the lack of genetic differentiation in neutral DNA. However, this appears to be insufficient not to classify them as species, as long as the calls can be regarded as diagnostic.     

Common Garden Experiment Reveals Genetic Control of Phenotypic Divergence between Swamp Sparrow Subspecies That Lack Divergence in Neutral Genotypes

Background

Adaptive divergence between populations in the face of strong selection on key traits can lead to morphological divergence between populations without concomitant divergence in neutral DNA. Thus, the practice of identifying genetically distinct populations based on divergence in neutral DNA may lead to a taxonomy that ignores evolutionarily important, rapidly evolving, locally-adapted populations. Providing evidence for a genetic basis of morphological divergence between rapidly evolving populations that lack divergence in selectively neutral DNA will not only inform conservation efforts but also provide insight into the mechanisms of the early processes of speciation. The coastal plain swamp sparrow, a recent colonist of tidal marsh habitat, differs from conspecific populations in a variety of phenotypic traits yet remains undifferentiated in neutral DNA.

Methods and Principal Findings

Here we use an experimental approach to demonstrate that phenotypic divergence between ecologically separated populations of swamp sparrows is the result of local adaptation despite the lack of divergence in neutral DNA. We find that morphological (bill size and plumage coloration) and life history (reproductive effort) differences observed between wild populations were maintained in laboratory raised individuals suggesting genetic divergence of fitness related traits.

Conclusions and Significance

Our results support the hypothesis that phenotypic divergence in swamps sparrows is the result of genetic differentiation, and demonstrate that adaptive traits have evolved more rapidly than neutral DNA in these ecologically divergent populations that may be in the early stages of speciation. Thus, identifying evolutionarily important populations based on divergence in selectively neutral DNA could miss an important level of biodiversity and mislead conservation efforts.     

Extrinsic and intrinsic factors influence fitness in an avian hybrid zone

The effects of hybridization on evolutionary processes are primarily determined by the differential between hybrid and parental species fitness. Assessing the impacts of hybridization can be challenging, however, as determining the relationship between individual fitness and the extent of introgression in wild populations is difficult. We evaluated the fitness consequences of hybridization for pure and hybrid females in a hybrid zone between two tidal marsh birds, the saltmarsh sparrow (Ammodramus caudacutus), a salt marsh obligate, and Nelson's sparrow (A. nelsoni), which has a broader ecological niche and a much younger evolutionary association with salt marshes. Biotic stressors associated with nesting in tidal environments suggest an important role for differential adaptation in shaping hybrid zone dynamics, with saltmarsh sparrows predicted to be better adapted to nesting in salt marshes. We collected DNA samples from adults (n = 394) and nestlings (n = 431) to determine the extent of introgression using 12 microsatellite loci and tested for the influence of extrinsic (nest placement) and intrinsic (genotype) factors on female reproductive success. We monitored nests (n = 228), collected data on reproductive output, and estimated daily nest survival rates using female genotype and nest characteristics as covariates. To test for reduced survival of hybrid females, we also used capture data to assess the distribution of admixed male and female individuals across age classes. Reproductive success of females varied by genotypic class, but hybrids did not have intermediate success as predicted. Instead, we found that pure Nelson's sparrows had, on average, 33% lower hatching success than any other genotype, whereas F1/F2 hybrids, backcrossed Nelson's sparrows, and backcrossed and pure saltmarsh sparrows all had similar hatching success. We found no effect of genotype or nest placement on daily nest survival probabilities. However, hybrid individuals with a higher proportion of saltmarsh sparrow alleles exhibit nesting behaviours better suited to nesting successfully in tidal marshes. Further, while the proportion of F1/F2 individuals was similar between nestling and adult males, we found that the proportion of F1/F2 individuals was 2.3 times greater in nestling females compared with adult females, indicating reduced survival of F1 females. We conclude that differences in reproductive success among pure and admixed individuals coupled with intrinsic mechanisms (reduced survival in F1 females) shape hybrid zone dynamics in this system.     

Habitat type and ambient temperature contribute to bill morphology

Avian bills are iconic structures for the study of ecology and evolution, with hypotheses about the morphological structure of bills dating back to Darwin. Several ecological and physiological hypotheses have been developed to explain the evolution of the morphology of bill shape. Here, we test some of these hypotheses such as the role of habitat, ambient temperature, body size, intraspecific competition, and ecological release on the evolution of bill morphology. Bill morphology and tarsus length were measured from museum specimens of yellow warblers, and grouped by habitat type, sex, and subspecies. We calculated the mean maximum daily temperature for the month of July, the hottest month for breeding specimens at each collecting location. Analysis of covariance models predicted total bill surface area as a function of sex, habitat type, body size, and temperature, and model selection techniques were used to select the best model. Habitat, mangrove forests compared with inland habitats, and climate had the largest effects on bill size. Coastal wetland habitats and island populations of yellow warblers had similar bill morphology, both of which are larger than mainland inland populations. Temperate but not tropical subspecies exhibited sexual dimorphism in bill morphology. Overall, this study provides evidence that multiple environmental factors, such as temperature and habitat, contribute to the evolution of bill morphology.     

Interspecific competition promotes habitat and morphological divergence in a secondary contact zone between two hybridizing songbirds

Interspecific competition is assumed to play an important role in the ecological differentiation of species and speciation. However, empirical evidence for competition's role in speciation remains surprisingly scarce. Here, we studied the role of interspecific competition in the ecological differentiation and speciation of two closely related songbird species, the Common Nightingale (Luscinia megarhynchos) and the Thrush Nightingale (Luscinia luscinia). Both species are insectivorous and ecologically very similar. They hybridize in a secondary contact zone, which is a mosaic of sites where both species co‐occur (syntopy) and sites where only one species is present (allotopy). We analysed fine‐scale habitat data for both species in both syntopic and allotopic sites and looked for associations between habitat use and bill morphology, which have been previously shown to be more divergent in sympatry than in allopatry. We found that the two nightingale species differ in habitat use in allotopic sites, where L. megarhynchos occurred in drier habitats and at slightly higher elevations, but not in syntopic sites. Birds from allotopic sites also showed higher interspecific divergence in relative bill size compared to birds from syntopic sites. Finally, we found an association between bill morphology and elevation. Our results are consistent with the view that interspecific competition in nightingales has resulted in partial habitat segregation in sympatry and that the habitat‐specific food supply has in turn very likely led to bill size divergence. Such ecological divergence may enhance prezygotic as well as extrinsic postzygotic isolation and thus accelerate the completion of the speciation process.     

Causes of reduced clutch size in a tidal marsh endemic

We tested three hypotheses of clutch size variation in two subspecies of the swamp sparrow (Melospiza georgiana georgiana and M. g. nigrescens). Swamp sparrows follow the pattern of other estuarine endemics, where clutch size is smaller among tidal salt marsh populations (M. g. nigrescens) than their closest inland relatives (M. g. georgiana). Our results support predation risk and temperature, but not adult survival, as explanations of this pattern in swamp sparrows. Coastal nests were twice as likely to fail as inland nests, and parental activity around the nest site was positively related to clutch size at both sites. When brood size was controlled for, coastal adults visited nests less often and females vocalized less frequently during visits than inland birds, which may decrease nest detectability to predators. Coastal parents waited longer than inland birds to feed offspring in the presence of a model nest predator, but there was no difference in their response to models of predators of adults, as would be expected if coastal birds possessed increased longevity. Additionally, coastal females laid more eggs than inland females over a single season, following a within-season bet-hedging strategy rather than reducing within-season investment. Coastal territories experienced ambient air temperatures above the physiological zero of egg development more often, and higher temperatures during laying correlated with smaller clutches and increased egg inviability among coastal birds. Similar effects were not seen among inland nests, where laying temperatures were generally below physiological zero. Both subspecies showed an increase in hatching asynchrony and a decrease in apparent incubation length under high temperatures. Coastal individuals, however, showed less hatching asynchrony overall despite higher temperatures. Both air temperatures during laying and predation risk could potentially explain reduced clutch size in not only coastal plain swamp sparrows, but also other tidal marsh endemics.     

Restoration of the Sieperda Tidal Marsh in the Scheldt Estuary, The Netherlands

Because of land reclamation, reinforcement of dikes, and the deepening of shipping channels, large areas of tidal marshes have been removed or eroded from the Scheldt estuary during the last two centuries. Tidal wetland restoration contributes toward compensating this loss of habitat. Not all restoration projects are meticulously planned, however; some are forced by nature. During a severe storm in 1990, a dike was breached in the brackish part of the Scheldt estuary and returned tidal influence to the Sieperda polder. In the 10 years since the dike breach, the former polder has changed into a brackish tidal marsh. Here we report on the geomorphologic and ecological developments that have taken place in the marsh. Tidal intrusion into the former polder turned crop fields into mudflats and changed pastures into salty marsh vegetation. The digging of a new creek improved marsh hydrology and enhanced tidal intrusion further into the marsh. Macrofauna typical of estuarine mudflats established rapidly in the developing marsh. Vegetation succession took place rapidly. Within 5 years, large areas of mudflats became covered with marsh vegetation. Birds characteristic of salt marshes were observed breeding or seen foraging in the marsh. The number of wading birds declined as areas of mudflat became overgrown. It is demonstrated that tidal flow is the engine to tidal marsh restoration. Tidal influence caused geomorphologic changes, which directed ecological developments in the former polder.     

The role of marker traits in the assortative mating within red crossbills, Loxia curvirostra complex

We conducted mate choice experiments to determine whether differences in calls or bill morphology might influence assortative mating between call types of red crossbills (Loxia curvirostra complex) that have diverged in bill structure to specialize on different species of conifers. Females preferred males that gave calls that matched their own type, but did not prefer males that more closely approximated the average or optimal bill size of the female's call type. These results were consistent with our breeding simulations, which showed that females gained an indirect fitness benefit by choosing a male of her own call type because this reduced the production of offspring with morphologies that fell between adaptive peaks. However, choice based on bill morphology within a call type provided no further benefit. Calls, which crossbills learn from their parents, likely act as a marker trait indicative of the morphological adaptations of the group, allow for easy assessment of potential mates and facilitate rapid divergence under ecological selection.     

Discordant patterns of genetic and phenotypic differentiation in five grasshopper species codistributed across a microreserve network

Conservation plans can be greatly improved when information on the evolutionary and demographic consequences of habitat fragmentation is available for several codistributed species. Here, we study spatial patterns of phenotypic and genetic variation among five grasshopper species that are codistributed across a network of microreserves but show remarkable differences in dispersal‐related morphology (body size and wing length), degree of habitat specialization and extent of fragmentation of their respective habitats in the study region. In particular, we tested the hypothesis that species with preferences for highly fragmented microhabitats show stronger genetic and phenotypic structure than codistributed generalist taxa inhabiting a continuous matrix of suitable habitat. We also hypothesized a higher resemblance of spatial patterns of genetic and phenotypic variability among species that have experienced a higher degree of habitat fragmentation due to their more similar responses to the parallel large‐scale destruction of their natural habitats. In partial agreement with our first hypothesis, we found that genetic structure, but not phenotypic differentiation, was higher in species linked to highly fragmented habitats. We did not find support for congruent patterns of phenotypic and genetic variability among any studied species, indicating that they show idiosyncratic evolutionary trajectories and distinctive demographic responses to habitat fragmentation across a common landscape. This suggests that conservation practices in networks of protected areas require detailed ecological and evolutionary information on target species to focus management efforts on those taxa that are more sensitive to the effects of habitat fragmentation.