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A genetic signature of the evolution of loss of flight in the Galapagos cormorant

INTRODUCTION Changes in the size and proportion of limbs and other structures have played a key role in the evolution of species. One common class of limb modification is recurrent wing reduction and loss of flight in birds. Indeed, Darwin used the occurrence of flightless birds as an argument in favor of his theory of natural selection. Loss of flight has evolved repeatedly and is found among 26 families of birds in 17 different orders. Despite the frequency of these modifications, we have a limited understanding of their underpinnings at the genetic and molecular levels. RATIONALE To better understand the evolution of changes in limb size, we studied a classic case of recent loss of flight in the Galapagos cormorant ( Phalacrocorax harrisi ). Cormorants are large water birds that live in coastal areas or near lakes, and P. harrisi is the only flightless cormorant among approximately 40 extant species. The entire population is distributed along the coastlines of Isabela and Fernandina islands in the Galapagos archipelago. P. harrisi has a pair of short wings, which are smaller than those of any other cormorant. The extreme reduction of the wings and pectoral skeleton observed in P. harrisi is an attractive model for studying the evolution of loss of flight because it occurred very recently; phylogenetic evidence suggests that P. harrisi diverged from its flighted relatives within the past 2 million years. We developed a comparative and predictive genomics approach that uses the genome sequences of P. harrisi and its flighted relatives to find candidate genetic variants that likely contributed to the evolution of loss of flight. RESULTS We sequenced and de novo assembled the whole genomes of P. harrisi and three closely related flighted cormorant species. We identified thousands of coding variants exclusive to P. harrisi and classified them according to their probability of altering protein function based on conservation. Variants most likely to alter protein function were significantly enriched in genes mutated in human skeletal ciliopathies, including Ofd1 , Evc , Wdr34 , and Ift122 . We carried out experiments in Caenorhabditis elegans to confirm that a missense variant present in the Galapagos cormorant IFT122 protein is sufficient to affect ciliary function. The primary cilium is essential for Hedgehog (Hh) signaling in vertebrates, and individuals affected by ciliopathies have small limbs and ribcages, mirroring the phenotype of P. harrisi . We also identified a 4–amino acid deletion in the regulatory domain of Cux1 , a highly conserved transcription factor that has been experimentally shown to regulate limb growth in chicken. The four missing amino acids are perfectly conserved in all birds and mammals sequenced to date. We tested the consequences of this deletion in a chondrogenic cell line and showed that it impairs the ability of CUX1 to transcriptionally up-regulate cilia-related genes (some of which contain function-altering variants in P. harrisi ) and to promote chondrogenic differentiation. Finally, we show that positive selection may have played a role in the fixation of the variants associated with loss of flight in P. harrisi . CONCLUSION Our results indicate that the combined effect of variants in genes necessary for the correct transcriptional regulation and function of the primary cilium likely contributed to the evolution of highly reduced wings and other skeletal adaptations associated with loss of flight in P. harrisi . Our approach may be generally useful for identification of variants underlying evolutionary novelty from genomes of closely related species.

Science↗

Genetic and phenotypic variation along an ecological gradient in lake trout Salvelinus namaycush

Background Adaptive radiation involving a colonizing phenotype that rapidly evolves into at least one other ecological variant, or ecotype, has been observed in a variety of freshwater fishes in post-glacial environments. However, few studies consider how phenotypic traits vary with regard to neutral genetic partitioning along ecological gradients. Here, we present the first detailed investigation of lake trout Salvelinus namaycush that considers variation as a cline rather than discriminatory among ecotypes. Genetic and phenotypic traits organized along common ecological gradients of water depth and geographic distance provide important insights into diversification processes in a lake with high levels of human disturbance from over-fishing. Results Four putative lake trout ecotypes could not be distinguished using population genetic methods, despite morphological differences. Neutral genetic partitioning in lake trout was stronger along a gradient of water depth, than by locality or ecotype. Contemporary genetic migration patterns were consistent with isolation-by-depth. Historical gene flow patterns indicated colonization from shallow to deep water. Comparison of phenotypic ( Pst ) and neutral genetic variation ( Fst ) revealed that morphological traits related to swimming performance (e.g., buoyancy, pelvic fin length) departed more strongly from neutral expectations along a depth gradient than craniofacial feeding traits. Elevated phenotypic variance with increasing water depth in pelvic fin length indicated possible ongoing character release and diversification. Finally, differences in early growth rate and asymptotic fish length across depth strata may be associated with limiting factors attributable to cold deep-water environments. Conclusion We provide evidence of reductions in gene flow and divergent natural selection associated with water depth in Lake Superior. Such information is relevant for documenting intraspecific biodiversity in the largest freshwater lake in the world for a species that recently lost considerable genetic diversity and is now in recovery. Unknown is whether observed patterns are a result of an early stage of incipient speciation, gene flow-selection equilibrium, or reverse speciation causing formerly divergent ecotypes to collapse into a single gene pool.

Lake Superior, Isle Royale↗

From top to bottom: Do Lake Trout diversify along a depth gradient in Great Bear Lake, NT, Canada?

Depth is usually considered the main driver of Lake Trout intraspecific diversity across lakes in North America. Given that Great Bear Lake is one of the largest and deepest freshwater systems in North America, we predicted that Lake Trout intraspecific diversity to be organized along a depth axis within this system. Thus, we investigated whether a deep-water morph of Lake Trout co-existed with four shallow-water morphs previously described in Great Bear Lake. Morphology, neutral genetic variation, isotopic niches, and life-history traits of Lake Trout across depths (0–150 m) were compared among morphs. Due to the propensity of Lake Trout with high levels of morphological diversity to occupy multiple habitat niches, a novel multivariate grouping method using a suite of composite variables was applied in addition to two other commonly used grouping methods to classify individuals. Depth alone did not explain Lake Trout diversity in Great Bear Lake; a distinct fifth deep-water morph was not found. Rather, Lake Trout diversity followed an ecological continuum, with some evidence for adaptation to local conditions in deep-water habitat. Overall, trout caught from deep-water showed low levels of genetic and phenotypic differentiation from shallow-water trout, and displayed higher lipid content (C:N ratio) and occupied a higher trophic level that suggested an potential increase of piscivory (including cannibalism) than the previously described four morphs. Why phenotypic divergence between shallow- and deep-water Lake Trout was low is unknown, especially when the potential for phenotypic variation should be high in deep and large Great Bear Lake. Given that variation in complexity of freshwater environments has dramatic consequences for divergence, variation in the complexity in Great Bear Lake (i.e., shallow being more complex than deep), may explain the observed dichotomy in the expression of intraspecific phenotypic diversity between shallow- vs. deep-water habitats. The ambiguity surrounding mechanisms driving divergence of Lake Trout in Great Bear Lake should be seen as reflective of the highly variable nature of ecological opportunity and divergent natural selection itself.

Northwest Territories↗

Genetic comparison of naturally spawned and artificially propagated Lake Ontario lake trout fry: Evaluation of a stocking strategy for species rehabilitation

Two strategies have been used in the effort to restore lake trout Salvelinus namaycush to Lake Ontario. First, lake trout strains from multiple wild and hatchery sources have been stocked to maximize genetic variability in the lake, Second, a unique hatchery “strain” of fish to be stocked was created each year with gametes collected from adult, hatchery‐origin fish that had survived to maturity after being stocked into Lake Ontario. Several hatchery strains may be represented among the adults captured and used to propagate this unique strain, termed the “Ontario strain.” The “Ontario strain” may have a genetically based potential for enhanced survival over other strains stocked in Lake Ontario because it is composed of the progeny of fish that have survived from the yearling to the adult life stage, Unlike naturally spawned fish, however, the “Ontario strain” has been shielded from natural selection during the critical period of mortality from spawning through the first year of life, The purpose of this study was to determine whether the “Ontario strain” was genetically representative of the wild fry produced in the lake, We examined the strain composition of three year‐classes of wild‐caught fry and six year‐classes of the “Ontario strain” using allozyme data with mixed‐stock analysis. The hatchery and wild fry were genetically dissimilar. In addition, the composition of the “Ontario strain” changed from predominantly Seneca strain in 1983–1984 to predominantly Superior and Killala strains from 1983 to 1989. Mixed‐stock estimates indicated that in contrast to the hatchery‐reared fry, strain composition of wildcaught fry did not vary greatly from year to year. Progeny of Seneca × Seneca crosses were the predominant fry in the three year‐classes of wild fry. The genetic dissimilarity between the “Ontario strain” fry and wild‐caught fry could be caused by differential mortality among the wild fry between spawning and fry emergence or by differential vulnerability of strains to the gill‐net sampling used to capture adults for gamete collection. Based on these results, the development of a new hatchery brood stock from wild‐caught fry is recommended as an alternative to the collection and propagation of gametes from mature hatchery‐origin lake trout. In addition, the composition of the hatchery strains stocked should be altered to emphasize those strains that reproduce successfully. Both options should be considered in order to speed the rehabilitation process.

Lake Ontario↗

Dispersal and survival of a polygynandrous passerine

Although sex biases in survival and dispersal are thought to be linked to avian mating systems, little is known about these demographic patterns in less common mating strategies such as polygynandry. We investigated breeding-site fidelity, natal philopatry, and apparent survival of the polygynandrous Smith's Longspur ( Calcarius pictus ) over a 7-yr period at 2 areas in Alaska's Brooks Range. We used capture–recapture histories of 243 color-banded adults and 431 juveniles to estimate annual survival and determined dispersal patterns from 34 adults that were found breeding within the study areas over multiple years. Most adults (88%) returned to nest in the same breeding neighborhood as in previous years; mean dispersal distance was 300.9 ± 74.2 m and did not differ between sexes. Juveniles exhibited low natal philopatry; only 4% of banded hatch-year birds were resighted as adults during subsequent years. Those that did return dispersed, on average, 1,674.4 ± 465.8 m from their natal nests ( n = 6). Model-averaged survival estimates indicated that annual survival of adult females (50–58%) was only slightly lower than that of males (60–63%); juvenile survival was 41% but was paired with a low (13%) encounter probability. We attribute the lack of sex bias in adult dispersal to this species' polygynandrous mating strategy. Within this system, there are multiple mates within a breeding neighborhood. We argue that natural selection may favor females that remain on the same, familiar breeding site, because they do not have to disperse to a new area to find a suitable mate. Dispersal among breeding populations most likely occurs by juveniles returning as adults. Our findings support hypotheses that suggest a relationship between dispersal and mating strategy and provide some of the first insight into the demographic patterns of a polygynandrous passerine.

The Auk↗

Foraging optimally for home ranges

Economic models predict behavior of animals based on the presumption that natural selection has shaped behaviors important to an animal's fitness to maximize benefits over costs. Economic analyses have shown that territories of animals are structured by trade-offs between benefits gained from resources and costs of defending them. Intuitively, home ranges should be similarly structured, but trade-offs are difficult to assess because there are no costs of defense, thus economic models of home-range behavior are rare. We present economic models that predict how home ranges can be efficient with respect to spatially distributed resources, discounted for travel costs, under 2 strategies of optimization, resource maximization and area minimization. We show how constraints such as competitors can influence structure of homes ranges through resource depression, ultimately structuring density of animals within a population and their distribution on a landscape. We present simulations based on these models to show how they can be generally predictive of home-range behavior and the mechanisms that structure the spatial distribution of animals. We also show how contiguous home ranges estimated statistically from location data can be misleading for animals that optimize home ranges on landscapes with patchily distributed resources. We conclude with a summary of how we applied our models to nonterritorial black bears (Ursus americanus) living in the mountains of North Carolina, where we found their home ranges were best predicted by an area-minimization strategy constrained by intraspecific competition within a social hierarchy. Economic models can provide strong inference about home-range behavior and the resources that structure home ranges by offering falsifiable, a priori hypotheses that can be tested with field observations.

Journal of Mammalogy↗

Comparative demography of new world populations of thrushes (Turdus spp.): Comment

Survival and fecundity are fundamental to the study of evolutionary ecology, as they are two of the key variables influencing the constrained optimization process we call natural selection. Likewise, population managers require accurate estimates of these parameters; along with dispersal, they govern population dynamics ( Lack 1954 ) and thus are essential for predicting population change and the effects of management actions. It can be frustrating, then, when survival is difficult to estimate for certain populations, as in some avian species that exhibit a substantial degree of dispersal.

Ecology↗

Structural determinations from diamond drilling

Several problems may arise in the geometrical determinations of strike and dip from core drilling. If the stratigraphic or structural surfaces are plane, as the bedding planes of a homocline, two problems may exist, depending upon whether the drill cores do or do not penetrate to some recognizable horizon. The first of these is solved by a simple graphic interpolation, and is not discussed in this paper. The second, however, is solved for the most general case of three non-parallel drill holes, no one of which is necessarily vertical. Errors are necessarily present in this solution, due to: 1. Inaccuracies in the measurement of the initial directions and dips of the drill holes. 2. Deviations of the drill holes from their initial directions. 3. Inaccuracies in the measurement of the angles between the axes of the drill cores, and the stratigraphic or structural planes. The second and third of these causes result in initial errors in the measurement of the core angles. Hence, for this simplest case, error formulm are developed for finding the final errors in the angles of strike and dip, under certain specified assumptions. To illustrate the solution, a problem is stated and solved. This problem is of a general nature, selected to show the necessary operations when the three drill holes occur in different octants. The error formulae are subsequently applied. For curved surfaces, as where folded beds are present, the same two problems arise, depending upon whether the drill holes do or do not penetrate to some recognizable horizon. Only the first of these problems is considered. It is shown, in this case, that 9 drill holes are required to obtain a satisfactory solution. The method consists essentially in fitting nine known points to a ternary quadric surface; and in determining the equations of the planes tangent to this surface at any of the nine points, or at any other points on the surface. The strikes and dips are then derived from these equations. © 1943 Society of Economic Geologists, Inc.

Economic Geology↗

The marine snail, Cerithidea californica, matures at smaller sizes where parasitism is high

I investigated life-history and parasitism in the salt marsh snail, Cerithidea californica. Latitude and growing conditions were important factors determining maturation size. After accounting for environmental variation, there was a negative association between the maturation size of snails and the prevalence of parasitic castration by larval trematodes. As predicted by life-history theory, this may represent an adaptation against parasitism that is similar to previous observations of life-history adaptations in species subject to predation or disturbance. However, it was unclear whether this adaptation was due to phenotypic plasticity or genetic differences among populations resulting from natural selection so I conducted a reciprocal transplant between sites with high and low prevalence and found source population differences in maturation size. It appears, therefore, that the life-history differences between these populations are at least partially genetic or may represent an adaptive developmental switch that was initiated prior to the transplant.

Oikos↗

Occurrence of volatile organic compounds in selected urban streams in the United States, 1995-2003

As part of the U.S. Geological Survey's (USGS) National Water-Quality Assessment (NAWQA) Program, urban indicator sites were monitored to (1) characterize the stream quality from drainage basins with predominantly residential and commercial land use, and (2) determine which selected natural and anthropogenic factors affect stream quality. A total of 869 water samples were collected from 37 urban streams during 1995-2003 and were analyzed for 87 volatile organic compounds (VOCs). The occurrence of VOCs in urban streams is described in this report for (1) all samples as a single dataset, (2) all samples grouped by streamflow pentiles, and (3) all samples grouped by warmer (April through September) and cooler (October through March) months by the detection frequency and (or) concentration of (a) any VOC, (b) VOC groups, and (c) individual compounds. An assessment level of 0.02 microgram per liter (ug/L) was used to compute the detection frequencies and concentrations of VOCs. Concentrations of VOCs were compared to (1) U.S. Environmental Protection Agency's (USEPA) drinking-water Maximum Contaminant Levels (MCLs) or Drinking Water Advisories, (2) Health-Based Screening Levels (HBSLs) developed by the USGS in collaboration with the USEPA and other agencies, and (3) USEPA and Canadian aquatic-life criteria. One or more VOCs were detected in 97.1 percent of 869 samples, and one or more VOCs were detected frequently (greater than 80 percent) at all sites. The median total VOC concentration for all samples was 0.57 ug/L, and total VOC concentrations in a single sample ranged from not detected to 698 ug/L. About 85 percent of the samples contained two or more VOCs, and about one-half contained five or more VOCs. The gasoline hydrocarbons were the most frequently occurring VOC group followed by solvents, trihalomethanes (THMs), gasoline oxygenates, organic synthesis compounds, fumigants, and refrigerants. Concentration ranges for most VOC groups were distributed over at least two orders of magnitude. Fifty-seven of the 87 VOCs analyzed were detected in at least one sample at an assessment level of 0.02 ug/L. More than one-half of the 30 VOCs not detected in samples were organic synthesis compounds. Fifteen compounds had detection frequencies greater than or equal to 10 percent. With the exception of toluene and chloroform, the median concentration of each VOC for all samples was less than the assessment level. Furthermore, the median concentrations of detections for the 15 most frequently occurring VOCs ranged from 0.03 to 3.9 ug/L, and typically were less than or equal to 0.10 ug/L. The 869 samples from the 37 sites were stratified into five streamflow pentiles (less than 20, 20-less than 40, 40-less than 60, 60-less than 80, and greater than or equal to 80 percent of estimated long-term streamflow statistics) for comparison of the occurrence of VOCs. The detection frequency of one or more VOCs by streamflow pentile varied only slightly from 96.7 to 97.7 percent. The median total VOC concentrations in samples for the five streamflow pentiles ranged from 0.39 to 1.0 ug/L. Two or more VOCs were present in more than 80 percent of samples in each of the five pentiles. The gasoline hydrocarbons, solvents, THMs, and gasoline oxygenates occurred frequently (greater than 30 percent) in all streamflow pentiles, in contrast to the organic synthesis compounds, fumigants, and refrigerants that occurred less frequently in urban streams under all streamflow conditions. The median total VOC concentrations for gasoline hydrocarbons, solvents, gasoline oxygenates, and organic synthesis compounds generally increased as streamflow increased. In contrast, the median total VOC concentrations for THMs and fumigants generally decreased as streamflow increased. The median total VOC concentrations for refrigerants showed no pattern as streamflow increased. Because differences between VOC occurrence and streamflow pentiles were small for most compariso

Scientific Investigations Report↗

Pesticides in Wyoming Groundwater, 2008-10

Groundwater samples were collected from 296 wells during 1995-2006 as part of a baseline study of pesticides in Wyoming groundwater. In 2009, a previous report summarized the results of the baseline sampling and the statistical evaluation of the occurrence of pesticides in relation to selected natural and anthropogenic (human-related) characteristics. During 2008-10, the U.S. Geological Survey, in cooperation with the Wyoming Department of Agriculture, resampled a subset (52) of the 296 wells sampled during 1995-2006 baseline study in order to compare detected compounds and respective concentrations between the two sampling periods and to evaluate the detections of new compounds. The 52 wells were distributed similarly to sites used in the 1995-2006 baseline study with respect to geographic area and land use within the geographic area of interest. Because of the use of different types of reporting levels and variability in reporting-level values during both the 1995-2006 baseline study and the 2008-10 resampling study, analytical results received from the laboratory were recensored. Two levels of recensoring were used to compare pesticides—a compound-specific assessment level (CSAL) that differed by compound and a common assessment level (CAL) of 0.07 microgram per liter. The recensoring techniques and values used for both studies, with the exception of the pesticide 2,4-D methyl ester, were the same. Twenty-eight different pesticides were detected in samples from the 52 wells during the 2008-10 resampling study. Pesticide concentrations were compared with several U.S. Environmental Protection Agency drinking-water standards or health advisories for finished (treated) water established under the Safe Drinking Water Act. All detected pesticides were measured at concentrations smaller than U.S. Environmental Protection Agency drinking-water standards or health advisories where applicable (many pesticides did not have standards or advisories). One or more pesticides were detected at concentrations greater than the CAL in water from 16 of 52 wells sampled (about 31 percent) during the resampling study. Detected pesticides were classified into one of six types: herbicides, herbicide degradates, insecticides, insecticide degradates, fungicides, or fungicide degradates. At least 95 percent of detected pesticides were classified as herbicides or herbicide degradates. The number of different pesticides detected in samples from the 52 wells was similar between the 1995-2006 baseline study (30 different pesticides) and 2008-2010 resampling study (28 different pesticides). Thirteen pesticides were detected during both studies. The change in the number of pesticides detected (without regard to which pesticide was detected) in groundwater samples from each of the 52 wells was evaluated and the number of pesticides detected in groundwater did not change for most of the wells (32). Of those that did have a difference between the two studies, 17 wells had more pesticide detections in groundwater during the 1995-2006 baseline study, whereas only 3 wells had more detections during the 2008-2010 resampling study. The difference in pesticide concentrations in groundwater samples from each of the 52 wells was determined. Few changes in concentration between the 1995-2006 baseline study and the 2008-2010 resampling study were seen for most detected pesticides. Seven pesticides had a greater concentration detected in the groundwater from the same well during the baseline sampling compared to the resampling study. Concentrations of prometon, which was detected in 17 wells, were greater in the baseline study sample compared to the resampling study sample from the same well 100 percent of the time. The change in the number of pesticides detected (without regard to which pesticide was detected) in groundwater samples from each of the 52 wells with respect to land use and geographic area was calculated. All wells with land use classified as agricultural had the same or a smaller number of pesticides detected in the resampling study compared to the baseline study. All wells in the Bighorn Basin geographic area also had the same or a smaller number of pesticides detected in the resampling study compared to the baseline study.

Wyoming↗

Statistical facilitation in environmental science: Integrating results from complementary statistical analyses can improve ecological interpretations

Professionals working in biological conservation seek to understand, manage, and restore populations of native organisms using many techniques. A common approach for this discipline is using long-term data collections to inform decision making. However, several quantitative issues complicate statistical analysis of monitoring datasets and can reduce the utility of results for conservation decision making. Integrating results from multiple analyses applied to the same dataset (i.e., approaching the same biological problem using different techniques) is one way to address concerns related to field data that violate statistical assumptions. This process allows data analysts, researchers, and managers to assemble insights based on the weight of evidence. Here we tested whether three different statistical techniques [(1) multiple logistic regression on original data, (2) multiple logistic regression on standardized data (i.e., mean of 0 and standard deviation of 1), and (3) random forest analysis] identified a similar hierarchy for selecting natural and anthropogenic habitat regressors. Our examination of how environmental variables affected Plains Minnow ( Hybognathus placitus ), a state-threatened fish, is relevant to other taxa and locations. We gained useful information from redundancies (i.e., agreements across analyses). New directions also emerged by addressing ambiguities (i.e., disagreements among results across analyses). When multiple analyses were integrated into one ecological story, a clearer interpretation emerged. Viewing different statistical tests as facilitators that provide mutual advantages can advance the understanding and application of statistical analyses applied to non-experimental field datasets.

Kansas↗

The role of genetics in chronic wasting disease of North American cervids

Chronic wasting disease (CWD) is a major concern for the management of North American cervid populations. This fatal prion disease has led to declines in populations which have high CWD prevalence and areas with both high and low infection rates have experienced economic losses in wildlife recreation and fears of potential spill-over into livestock or humans. Research from human and veterinary medicine has established that the prion protein gene ( Prnp ) encodes the protein responsible for transmissible spongiform encephalopathies (TSEs). Polymorphisms in the Prnp gene can lead to different prion forms that moderate individual susceptibility to and progression of TSE infection. Prnp genes have been sequenced in a number of cervid species including those currently infected by CWD (elk, mule deer, white-tailed deer, moose) and those for which susceptibility is not yet determined (caribou, fallow deer, sika deer). Over thousands of sequences examined, the Prnp gene is remarkably conserved within the family Cervidae; only 16 amino acid polymorphisms have been reported within the 256 amino acid open reading frame in the third exon of the Prnp gene. Some of these polymorphisms have been associated with lower rates of CWD infection and slower progression of clinical CWD. Here we review the body of research on Prnp genetics of North American cervids. Specifically, we focus on known polymorphisms in the Prnp gene, observed genotypic differences in CWD infection rates and clinical progression, mechanisms for genetic TSE resistance related to both the cervid host and the prion agent and potential for natural selection for CWD-resistance. We also identify gaps in our knowledge that require future research.

Prion↗

Development of migratory behavior in northern white-tailed deer

I examined the development of migratory behavior in northern white-tailed deer (Odocoileus virginianus) from 1975 to 1996 by radio-tracking adult females and their fawns. Of 40 migratory fawns with radio-collared mothers, all returned from winter ranges to their mothers' summer ranges, as did 36 fawns with unknown mothers. Of 1.5- to 3.0-year-old daughters with radio-collared mothers, 67-80% continued migrating with mothers to their traditional summer ranges. Eighty-four percent (16/19) of yearling dispersers continued migratory behavior after replacing their natal summer ranges with their dispersal ranges, and 88% (14/16) of these continued migrating to their natal winter ranges, some through at least 6.5 years of age. Twenty percent (4/20) of nonmigratory fawns dispersed as yearlings, and two became migratory between their dispersal summer ranges and new winter ranges, one through 4.9 years of age and another through 6.5 years. Seven fawns changed their movement behavior from migratory to nonmigratory or vice versa as yearlings or when older, indicating that migratory behavior is not under rigid genetic control. Thus, the adaptiveness of migration must depend upon natural selection operating upon varying capacities and propensities to learn and mimic long-distance movements and not upon migratory behavior directly.

Canadian Journal of Zoology↗

Heterozygosity and fitness: No strong association in Great Lakes populations of the zebra mussel, Dreissena Polymorpha (Pallas)

A number of studies have found positive associations between allozyme heterozygosity and fitness surrogates (e.g., body size and growth rate) for marine molluscs. We investigated whether similar relationships exist for freshwater populations of the zebra mussel, Dreissena polymorpha. Only one significant correlation between multi-locus heterozygosity and shell length was observed for a total of 22 D. polymorpha populations surveyed from midwestern U.S.A. lakes and streams, and the result was not significant on a table-wide basis. Meta-analysis revealed a significant common correlation coefficient (effect magnitude) between multi-locus heterozygosity and shell length across all 22 sites (rc = 0.052, P = 0.019, 1557 df). However, the variance in shell length explained by multi-locus heterozygosity was small (rc2 = 0.0027), implying a weak causal relationship if any. Also, we saw no relationship between heterozygosity and growth rate in a one-year field enclosure experiment. A significant heterozygosity-shell length correlation previously reported for a zebra mussel population at Put-in-Bay, Lake Erie, Ohio, may have been the product of unique population dynamics, rather than natural selection. Similar demographic considerations may contribute to inconsistencies in heterozygosity-fitness correlations seen for other molluscs.

Malacologia↗

Mangrove removal in the belize cays: effects on mangrove-associated fish assemblages in the intertidal and subtidal

We investigated the effects of mangrove cutting on fish assemblages in Twin Cays, Belize, in two habitat types. We conducted visual censuses at two sites in adjoining undisturbed/disturbed (30%–70% of shoreline fringe removed) sub-tidal fringing Rhizophora mangle Linnaeus, 1753. Observers recorded significantly more species and individuals in undisturbed sites, especially among smaller, schooling species (e.g., atherinids, clupeids), where densities were up to 200 times greater in undisturbed habitat. Multivariate analyses showed distinct species assemblages between habitats at both sites. In addition, extensive trapping with wire minnow traps within the intertidal zone in both undisturbed and disturbed fringing and transition (landward) mangrove forests was conducted. Catch rates were low: 638 individuals from 24 species over 563 trap-nights. Trap data, however, indicated that mangrove disturbance had minimal effect on species composition in either forest type (fringe/transition). Different results from the two methods (and habitat types) may be explained by two factors: (1) a larger and more detectable species pool in the subtidal habitat, with visual "access" to all species, and (2) the selective nature of trapping. Our data indicate that even partial clearing of shoreline and more landward mangroves can have a significant impact on local fish assemblages.

Twin Cays↗

Genetic differences between hatchery and wild steelhead for growth and survival in the hatchery and seaward migration after release (Study sites: Dworshak Hatchery and Clearwater Hatchery; Stocks: Dworshak hatchery and Selway River wild; Year classes: 1994 and 1995)

Various studies suggest that sea ranching of anadromous salmonids can result in domestication (increased fitness in the hatchery program) and a loss of fitness for natural production; however, the mechanism has not been characterized adequately. We artificially spawned hatchery and wild steelhead Oncorhynchus mykiss from the Clearwater River, Idaho, reared the resulting genetically marked (at the PEPA allozyme locus) progeny (HxH, HxW from hatchery females and wild males, and WxW) in hatcheries, and tested for differences in survival, growth, early maturation, downstream migration, and adult returns. Rearing treatments were mixed (crosses reared together) and separate (crosses reared separately from each other) at the hatchery of origin for the hatchery population where smolts are produced in one year, and at a nearby hatchery employing lower rations, lower winter temperatures, and two years of rearing to more closely mimic the natural life history (natural smolt age = 2-4 years). The hatchery population had been artificially propagated for six generations at the onset of our study. We found little or no difference in survival in the hatchery but substantially higher rates of growth and subsequent downstream migration for HxH than for WxW fish. Faster growth for HxH fish resulted in greater size at release which contributed to their higher migration rate, but other as yet uncharacterized traits also affected migration since the migration difference between crosses was apparent even within size classes. Growth of WxW fish was slower in the mixed than in the separate treatment indicating that WxW fish were competitively inferior to HxH fish in the hatchery environment. Incidence of precocious males was higher for WxW than for HxH fish in the separate but not in the mixed treatment. Incidence of HxH precocious males was similar between treatments. Apparently, the presence of HxH fish suppressed high incidence of early maturation by WxW males. A direct effect beyond the suppression of WxW growth by HxH fish was involved because the effect persisted within size categories. In-hatchery survival and growth of WxW relative to HxH fish may have been better with two-year rearing than in the standard one-year program (differences were consistent but small and non-significant); however, performance remained substantially worse for progeny of wild fish. Greater downstream migration for HxH than for WxW fish was primarily due to greater residualization for WxW than for HxH fish near the smolt release site rather than to immediate differential mortality. By August the residuals had lost condition compared to their condition in the hatchery the previous March, a month before release, and the residuals produced almost no smolts the following spring. Adult return rate was higher for HxH than for WxW fish for one year-class, consistent with the difference in downstream migration; only three adults (all WxW) returned from the other. Intermediate performance by HxW fish on growth, early maturation, downstream migration, and adult returns corroborated the genetic basis of the stock differences. Natural selection after release from the hatchery favored fish that performed well in the hatchery (grew fast, didn’t mature early, and excelled in other as yet uncharacterized traits) and genetically changed (domesticated) the wild population to resemble the hatchery population.

Idaho↗

Genetic and environmental influences on cold hardiness of native and introduced riparian trees

To explore latitudinal genetic variation in cold hardiness and leaf phenology, we planted a common garden of paired collections of native and introduced riparian trees sampled along a latitudinal gradient. The garden in Fort Collins, Colorado (latitude 40.6°N), included 681 native plains cottonwood (Populus deltoides subsp. monilifera) and introduced saltcedar (Tamarix ramosissima, T. chinensis, and hybrids) collected from 15 sites from 29.2 to 47.6°N in the central United States. In the common garden, both species showed latitudinal variation in fall, but not spring, leaf phenology. This suggests that latitudinal gradient field observations in fall phenology are a result, at least in part, of the inherited variation in the critical photoperiod. Conversely, the latitudinal gradient field observations in spring phenology are largely a plastic response to the temperature gradient. Populations from higher latitudes exhibited earlier bud set and leaf senescence. Cold hardiness varied latitudinally in both fall and spring for both species. Although cottonwood was hardier than saltcedar in midwinter, the reverse was true in late fall and early spring. The latitudinal variation in fall phenology and cold hardiness of saltcedar appears to have developed as a result of multiple introductions of genetically distinct populations, hybridization, and natural selection in the 150 years since introduction.

Conference Paper↗