The fitness landscape of a community of Darwin's finches.

Divergent natural selection should lead to adaptive radiation-that is, the rapid evolution of phenotypic and ecological diversity originating from a single clade. The drivers of adaptive radiation have often been conceptualized through the concept of "adaptive landscapes," yet formal empirical estimates of adaptive landscapes for natural adaptive radiations have proven elusive. Here, we use a 17-year dataset of Darwin's ground finches (Geospiza spp.) at an intensively studied site on Santa Cruz (Galápagos) to estimate individual apparent lifespan in relation to beak traits. We use these estimates to model a multi-species fitness landscape, which we also convert to a formal adaptive landscape. We then assess the correspondence between estimated fitness peaks and observed phenotypes for each of five phenotypic modes (G. fuliginosa, G. fortis [small and large morphotypes], G. magnirostris, and G. scandens). The fitness and adaptive landscapes show 5 and 4 peaks, respectively, and, as expected, the adaptive landscape was smoother than the fitness landscape. Each of the five phenotypic modes appeared reasonably close to the corresponding fitness peak, yet interesting deviations were also documented and examined. By estimating adaptive landscapes in an ongoing adaptive radiation, our study demonstrates their utility as a quantitative tool for exploring and predicting adaptive radiation.

Grooming Time Predicts Survival: American Kestrels, Falco sparverius, on a Subtropical Island.

AbstractAnimals have evolved a variety of adaptations to care for their body surfaces, such as grooming behavior, which keeps the integument clean, parasite-free, and properly arranged. Despite extensive research on the grooming of mammals, birds, and arthropods, the survival value of grooming has never been directly measured in natural populations. We monitored grooming and survival in a population of marked American kestrels (Falco sparverius) on San Salvador Island, Bahamas. We found a strong association between time spent grooming and survival over a 2-year period. The quadratic relationship we show is consistent with stabilizing natural selection on grooming time. To our knowledge, this is the first evidence for a correlation between grooming time and survival in a natural population. Grooming time may predict the survival of many animal taxa, but additional studies are needed to determine the shape and strength of the relationship among birds, mammals, and arthropods.

The High-Elevation Peatlands of the Northern Andes, Colombia.

Andean peatlands are important carbon reservoirs for countries in the northern Andes and have a unique diversity. Peatland plant diversity is generally related to hydrology and water chemistry, and the response of the vegetation in tropical high-elevation peatlands to changes in elevation, climate, and disturbance is poorly understood. Here, we address the questions of what the main vegetation types of peat-forming vegetation in the northern Andes are, and how the different vegetation types are related to water chemistry and pH. We measured plant diversity in 121 peatlands. We identified a total of 264 species, including 124 bryophytes and 140 vascular plants. We differentiated five main vegetation types: cushion plants, Sphagnum, true mosses, sedges, and grasses. Cushion-dominated peatlands are restricted to elevations above 4000 m. Variation in peatland vegetation is mostly driven be elevation and water chemistry. Encroachment of sedges and Sphagnum sancto-josephense in disturbed sites was associated with a reduction in soil carbon. We conclude that peatland variation is driven first by elevation and climate followed by water chemistry and human disturbances. Sites with higher human disturbances had lower carbon content. Peat-forming vegetation in the northern Andes was unique to each site bringing challenges on how to better conserve them and the ecosystem services they offer.

Temporally varying disruptive selection in the medium ground finch (Geospiza fortis).

Disruptive natural selection within populations exploiting different resources is considered to be a major driver of adaptive radiation and the production of biodiversity. Fitness functions, which describe the relationships between trait variation and fitness, can help to illuminate how this disruptive selection leads to population differentiation. However, a single fitness function represents only a particular selection regime over a single specified time period (often a single season or a year), and therefore might not capture longer-term dynamics. Here, we build a series of annual fitness functions that quantify the relationships between phenotype and apparent survival. These functions are based on a 9-year mark-recapture dataset of over 600 medium ground finches (Geospiza fortis) within a population bimodal for beak size. We then relate changes in the shape of these functions to climate variables. We find that disruptive selection between small and large beak morphotypes, as reported previously for 2 years, is present throughout the study period, but that the intensity of this selection varies in association with the harshness of environment. In particular, we find that disruptive selection was strongest when precipitation was high during the dry season of the previous year. Our results shed light on climatic factors associated with disruptive selection in Darwin's finches, and highlight the role of temporally varying fitness functions in modulating the extent of population differentiation.

Alien Invasion: Biology of Philornis Flies Highlighting Philornis downsi, an Introduced Parasite of Galápagos Birds.

The muscid genus Philornis comprises approximately 50 described species of flies, nearly all of which are obligate parasites of nestling birds. Philornis species are native to the Neotropics and widely distributed from Florida to Argentina. Most research on this group has focused on P. downsi, which was introduced to the Galápagos Islands in the late twentieth century. Although Philornis parasitism kills nestlings in several native host species, nowhere do the effects seem more severe than in P. downsi in the Galápagos. Here, we review studies of native and introduced Philornis in an attempt to identify factors that may influence virulence and consider implications for the conservation of hosts in the Galápagos.

Epigenetic variation between urban and rural populations of Darwin's finches.

BACKGROUND: The molecular basis of evolutionary change is assumed to be genetic variation. However, growing evidence suggests that epigenetic mechanisms, such as DNA methylation, may also be involved in rapid adaptation to new environments. An important first step in evaluating this hypothesis is to test for the presence of epigenetic variation between natural populations living under different environmental conditions. RESULTS: In the current study we explored variation between populations of Darwin's finches, which comprise one of the best-studied examples of adaptive radiation. We tested for morphological, genetic, and epigenetic differences between adjacent "urban" and "rural" populations of each of two species of ground finches, Geospiza fortis and G. fuliginosa, on Santa Cruz Island in the Galápagos. Using data collected from more than 1000 birds, we found significant morphological differences between populations of G. fortis, but not G. fuliginosa. We did not find large size copy number variation (CNV) genetic differences between populations of either species. However, other genetic variants were not investigated. In contrast, we did find dramatic epigenetic differences between the urban and rural populations of both species, based on DNA methylation analysis. We explored genomic features and gene associations of the differentially DNA methylated regions (DMR), as well as their possible functional significance. CONCLUSIONS: In summary, our study documents local population epigenetic variation within each of two species of Darwin's finches.

Galápagos mockingbirds tolerate introduced parasites that affect Darwin's finches.

Introduced parasites threaten native host species that lack effective defenses. Such parasites increase the risk of extinction, particularly in small host populations like those on islands. If some host species are tolerant to introduced parasites, this could amplify the risk of the parasite to vulnerable host species. Recently, the introduced parasitic nest fly Philornis downsi has been implicated in the decline of Darwin's finch populations in the Galápagos Islands. In some years, 100% of finch nests fail due to P. downsi; however, other common host species nesting near Darwin's finches, such as the endemic Galápagos mockingbird (Mimus parvulus), appear to be less affected by P. downsi. We compared effects of P. downsi on mockingbirds and medium ground finches (Geospiza fortis) on Santa Cruz Island in the Galápagos. We experimentally manipulated the abundance of P. downsi in nests of mockingbirds and finches to measure the direct effect of the parasite on the reproductive success of each species of host. We also compared immunological and behavioral responses by each species of host to the fly. Although nests of the two host species had similar parasite densities, flies decreased the fitness of finches but not mockingbirds. Neither host species had a significant antibody-mediated immune response to P. downsi. Moreover, finches showed no significant increase in begging, parental provisioning, or plasma glucose levels in response to the flies. In contrast, parasitized mockingbird nestlings begged more than nonparasitized mockingbird nestlings. Greater begging was correlated with increased parental provisioning behavior, which appeared to compensate for parasite damage. The results of our study suggest that finches are negatively affected by P. downsi because they do not have such behavioral mechanisms for energy compensation. In contrast, mockingbirds are capable of compensation, making them tolerant hosts, and a possible indirect threat to Darwin's finches.

Galápagos mockingbirds tolerate introduced parasites that affect Darwin's finches.

Introduced parasites threaten native host species that lack effective defenses. Such parasites increase the risk of extinction, particularly in small host populations like those on islands. If some host species are tolerant to introduced parasites, this could amplify the risk of the parasite to vulnerable host species. Recently, the introduced parasitic nest fly Philornis downsi has been implicated in the decline of Darwin's finch populations in the Galápagos Islands. In some years, 100% of finch nests fail due to P. downsi; however, other common host species nesting near Darwin's finches, such as the endemic Galápagos mockingbird (Mimus parvulus), appear to be less affected by P. downsi. We compared effects of P. downsi on mockingbirds and medium ground finches (Geospiza fortis) on Santa Cruz Island in the Galápagos. We experimentally manipulated the abundance of P. downsi in nests of mockingbirds and finches to measure the direct effect of the parasite on the reproductive success of each species of host. We also compared immunological and behavioral responses by each species of host to the fly. Although nests of the two host species had similar parasite densities, flies decreased the fitness of finches but not mockingbirds. Neither host species had a significant antibody-mediated immune response to P. downsi. Moreover, finches showed no significant increase in begging, parental provisioning, or plasma glucose levels in response to the flies. In contrast, parasitized mockingbird nestlings begged more than nonparasitized mockingbird nestlings. Greater begging was correlated with increased parental provisioning behavior, which appeared to compensate for parasite damage. The results of our study suggest that finches are negatively affected by P. downsi because they do not have such behavioral mechanisms for energy compensation. In contrast, mockingbirds are capable of compensation, making them tolerant hosts, and a possible indirect threat to Darwin's finches.

Darwin's finches combat introduced nest parasites with fumigated cotton.

Introduced parasites are a threat to biodiversity when naïve hosts lack effective defenses against such parasites [1]. Several parasites have recently colonized the Galápagos Islands, threatening native bird populations [2]. For example, the introduced parasitic nest fly Philornis downsi (Diptera: Muscidae) has been implicated in the decline of endangered species of Darwin's finches, such as the mangrove finch (Camarhynchus heliobates) [3]. Here, we show that Darwin's finches can be encouraged to 'self-fumigate' nests with cotton fibers that have been treated with permethrin. Nests with permethrin-treated cotton had significantly fewer P. downsi than control nests, and nests containing at least one gram of cotton were virtually parasite-free. Nests directly fumigated with permethrin had fewer parasites and fledged more offspring than nests treated with water.

Diversity of feather mites (Acari: Astigmata) on Darwin's finches.

Feather mites are a diverse group of ectosymbionts that occur on most species of birds. Although Darwin's finches are a well-studied group of birds, relatively little is known about their feather mites. Nearly 200 birds across 9 finch species, and from 2 locations on Santa Cruz Island, Galápagos, were dust-ruffled during the 2009 breeding season. We found 8 genera of feather mites; the most prevalent genus was Mesalgoides (53-55%), followed by Trouessartia (40-45%), Amerodectes and Proctophyllodes (26-33%), Xolalgoides (21-27%), Analges and Strelkoviacarus (0-6%), and Dermoglyphus (2-4%). There was no evidence for microclimatic effects (ambient temperature and relative humidity) on mite diversity. Host body mass was significantly correlated with mean feather mite abundance across 7 of 8 well-sampled species of finches. Certhidea olivacea, the smallest species, did not fit this pattern and had a disproportionately high number of mites for its body mass.

Localized thermal anomalies in haloclines of coastal Yucatan sinkholes.

A temperature spike is reported in the haloclines of three Yucatan sinkholes along a 1 km NW-SE transect from 5 to 4 km inland from the Caribbean coast. The temperature spike decreases in magnitude from 3.5 degrees C to 0.2 degrees C, approaching the coast. The anomaly does not vary diurnally and does not extend down into the underlying sea water. These conditions are inconsistent with explanations such as radiation absorption within the halocline, in situ microbially mediated sulfate reduction within the halocline and the underlying sea water, and sulfide oxidation by photosynthetic purple and green bacteria within the halocline. One explanation consistent with the shape and halocline location of the temperature spike involves a localized sea water convection cell operating near the coast. Cold sea water from the Caribbean Sea enters the coastal limestone at depths of a few hundred meters and heats up because of the geothermal gradient, buoyantly rising in vertical fractures within the unconfined aquifer. Blocked by the less dense fresh water, the movement stops in the halocline where the warm sea water mixes with brackish water. The convection cycle would be completed by the coastward movement of cooling brackish water. The observed temperature anomalies could possibly be snapshots of this warm layer moving coastward.

The population genetics of host specificity: genetic differentiation in dove lice (Insecta: Phthiraptera).

Some species of parasites occur on a wide range of hosts while others are restricted to one or a few host species. The host specificity of a parasite species is determined, in part, by its ability to disperse between host species. Dispersal limitations can be studied by exploring the genetic structure of parasite populations both within a single species of host and across multiple host species. In this study we examined the genetic structure in the mitochondrial cytochrome oxidase I (COI) gene of two genera of lice (Insecta: Phthiraptera) occurring on multiple sympatric species of doves in southern North and Central America. One genus, Columbicola, is generally less host-specific than the other, Physconelloides. For both genera we identified substantial genetic differentiation between populations of conspecific lice on different host species, generally 10-20% sequence divergence. This level of divergence is in the range of that often observed between species of these two genera. We used nested clade analysis to explore fine scale genetic structure within species of these feather lice. We found that species of Physconelloides exhibited more genetic structure, both among hosts and among geographical localities, than did species of Columbicola. In many cases, single haplotypes within species of Columbicola are distributed on multiple host species. Thus, the population genetic structure of species of Physconelloides reveals evidence of geographical differentiation on top of high host species specificity. Underlying differences in dispersal biology probably explain the differences in population genetic structure that we observed between Columbicola and Physconelloides.