Decomposing the mid-Holocene Tsuga decline in eastern North America

The mid-Holocene decline of Tsuga canadensis (hereafter Tsuga) populations across eastern North America is widely perceived as a synchronous event, driven by pests/pathogens, rapid climate change, or both. Pattern identification and causal attribution are hampered by low stratigraphic density of pollen-sampling and radiometric dates at most sites, and by absence of highly resolved, paired pollen and paleoclimate records from single sediment cores, where chronological order of climatic and vegetational changes can be assessed. We present an intensely sampled (contiguous 1-cm intervals) record of pollen and water table depth (inferred from testate amoebae) from a single core spanning the Tsuga decline at Irwin Smith Bog in Lower Michigan, with high-precision chronology. We also present an intensively sampled pollen record from Tower Lake in Upper Michigan. Both sites show high-magnitude fluctuations in Tsuga pollen percentages during the pre-decline maximum. The terminal decline is dated at both sites ca. 5000 cal yr BP, some 400 years later than estimates from other sites and data compilations. The terminal Tsuga decline was evidently heterochronous across its range. A transient decline ca. 5350 cal yr BP at both sites may correspond to the terminal decline at other sites in eastern North America. At Irwin Smith Bog, the terminal Tsuga decline preceded an abrupt and persistent decline in water table depths by approximately 200 years, suggesting the decline was not directly driven by abrupt climate change. The Tsuga decline may best be viewed as comprising at least three phases: a long-duration pre-decline maximum with high-magnitude and high-frequency fluctuations, followed by a terminal decline at individual sites, followed in turn by two millennia of persistently low Tsuga populations. These phases may not be causally linked, and may represent dynamics taking place at multiple temporal and spatial scales. Further progress toward understanding the phenomenon requires an expanded network of high-resolution pollen and paleoclimate chronologies.     

Monarch butterfly migration and parasite transmission in eastern North America

Seasonal migration occurs in many animal systems and is likely to influence interactions between animals and their parasites. Here, we focus on monarch butterflies (Danaus plexippus) and a protozoan parasite (Ophryocystis elektroscirrha) to investigate how host migration affects infectious disease processes. Previous work showed that parasite prevalence was lower among migratory than nonmigratory monarch populations; two explanations for this pattern are that (1) migration allows animals to periodically escape contaminated habitats (i.e., migratory escape), and (2) long-distance migration weeds out infected animals (i.e., migratory culling). We combined field-sampling and analysis of citizen science data to examine spatiotemporal trends of parasite prevalence and evaluate evidence for these two mechanisms. Analysis of within-breeding-season variation in eastern North America showed that parasite prevalence increased from early to late in the breeding season, consistent with the hypothesis of migratory escape. Prevalence was also positively related to monarch breeding activity, as indexed by larval density. Among adult monarchs captured at different points along the east coast fall migratory flyway, parasite prevalence declined as monarchs progressed southward, consistent with the hypothesis of migratory culling. Parasite prevalence was also lower among monarchs sampled at two overwintering sites in Mexico than among monarchs sampled during the summer breeding period. Collectively, these results indicate that seasonal migration can affect parasite transmission in wild animal populations, with implications for predicting disease risks for species with threatened migrations.     

Genetic studies of laboratory-rearedMytilus edulis

Laboratory cultures of offspring from single families and a mass mating ofMytilus edulis were reared to the spat stage at different temperatures. Some cultures were reared further to the juvenile stage at the same, or at an altered temperature. Using electrophoresis, frequencies of genotypes at theOdh, Pgm, Gpi, Hex, EsD, Pgk andDia loci were recorded at the spat stage and, again, where possible, in juveniles. Significant deviations from expected genotype frequencies were observed at most loci in at least some cultures, but in no cases were these deviations related to the temperature at which the cultures had been reared. We speculate that these differences may have been due to variations in the density of mussel spat in the cultures. Very extensive and consistentPgm genotype-specific mortalities were evident in one family, and, in another family, consistent deviations were seen at theHex locus. Weaker, but nevertheless significant, deviations from expected genotype frequencies occurred at most other loci and it is concluded that, in most cases, the loci were acting as markers for a locus, or loci, on the same chromosome which had a strong effect on fitness. It is thought unlikely that evidence for similar levels of selection would be found in natural populations because, in wild populations, particular marker-locus genotypes would be associated with a range of genetic backgrounds rather than a single chromosome.     

Modeling range dynamics in heterogeneous landscapes: invasion of the hemlock woolly adelgid in eastern North America

Range expansion by native and exotic species will continue to be a major component of global change. Anticipating the potential effects of changes in species distributions requires models capable of forecasting population spread across realistic, heterogeneous landscapes and subject to spatiotemporal variability in habitat suitability. Several decades of theory and model development, as well as increased computing power and availability of fine-resolution GIS data, now make such models possible. Still unanswered, however, is the question of how well this new generation of dynamic models will anticipate range expansion. Here we develop a spatially explicit stochastic model that combines dynamic dispersal and population processes with fine-resolution maps characterizing spatiotemporal heterogeneity in climate and habitat to model range expansion of the hemlock woolly adelgid (HWA; Adelges tsugae). We parameterize this model using multiyear data sets describing population and dispersal dynamics of HWA and apply it to eastern North America over a 57-year period (1951-2008). To evaluate the model, the observed pattern of spread of HWA during this same period was compared to model predictions. Our model predicts considerable heterogeneity in the risk of HWA invasion across space and through time, and it suggests that spatiotemporal variation in winter temperature, rather than hemlock abundance, exerts a primary control on the spread of HWA. Although the simulations generally matched the observed current extent of the invasion of HWA and patterns of anisotropic spread, it did not correctly predict when HWA was observed to arrive in different geographic regions. We attribute differences between the modeled and observed dynamics to an inability to capture the timing and direction of long-distance dispersal events that substantially affected the ensuing pattern of spread.     

Morphological variation of Mytilus edulis and Mytilus trossulus in eastern Newfoundland

Allopatric populations of Mytilus species show distinct shell morphology which may be due to genetic and/or environmental effects. Sympatric populations of Mytilus species show similar shell morphology which may be due to hybridization eroding morphological differences and/or the influence of common environmental conditions. The present study examined shell morphology and shell shape from 16 sites in eastern Newfoundland where M. edulis L. and M. trossulus Gould coexist in common environments with limited hybridization. Shell morphology was based on measurements of eight characters, and shell shape was quantified by elliptic Fourier analysis of shell outlines. Significant differences were observed between species for both shell morphology and shell shape across 16 sites sampled. The relatively small differences in morphology and shape between the species were probably due to exposure to common environments rather than hybridization. Shell shape for M. edulis was more eccentric compared to M. trossulus which was more elongated. Shell shape analysis of a range of size classes at one site showed a change from an eccentric to an elongated shape going from the smaller to the larger size classes. Both species showed a similar trend, with the larger M. edulis more eccentric and the larger M. trossulus more elongated. Received: 17 July 1998 / Accepted: 6 January 1999     

Frequency, not relative abundance, of temperate tree species varies along climate gradients in eastern North America

There have been many attempts to model the impacts of climate change on the distributions of temperate tree species, but empirical analyses of the effects of climate on the distribution and abundance of tree species have lagged far behind the models. Here, we used forest inventory data to characterize variation in adult tree abundance along climate gradients for the 24 most common tree species in the northeastern United States. The two components of our measure of species abundance--local frequency vs. relative abundance--showed dramatically different patterns of variation along gradients of mean annual temperature and precipitation. Local frequency (i.e., the percentage of plots in a given climate in which a species occurred) varied strongly for all 24 species, particularly as a function of temperature. Relative abundance when present in a plot, on the other hand, was effectively constant for most species right up to their estimated climatic range limits. Although the range limits for both temperature and precipitation were quite broad for all of the species, the range of climates within which a species was common (i.e., high frequency) was much narrower. Because frequency in sites within a given climate shows a strong sensitivity to temperature, at least, this suggests that the processes determining canopy tree recruitment on new sites also vary strongly with climate.     

Genetic and nutritional control of the gametogenic cycle in Mytilus edulis

Individuals of Mytilus edulis were collected from two populations in easterm Long Island Sound, USA, at 6 wk intervals from January 1982 to February 1983, and the hypothesis that nutrient availability controls the timing of gametogenic events in this mussel was tested using genetic variants at the Lap locus. In eastern Long Island Sound, Lap genotypes differ in the net rate of nitrogen accumulation. A highly significant difference in reproductive cycle among genotypes occurred in a population of M. edulis where Lap genotypes differ in nitrogen budget. Genotypes with lower rates of nutrient accumulation delayed the reproductive cycle by approximately six weeks relative to genotypes with high rates of nutrient accumulation. No significant difference in reproductive cycle occurred at a locality where Lap genotypes do not differ in nitrogen budget. These data indicate that nutritional status can exert significant control over the timing of gametogenesis in M. edulis. Differences in reproductive cycle among Lap genotypes provide a mechanism of non-random mating among genotypes which may result in heterozygote deficiencies at this locus.     

Evolutionary genetics of the Mytilus edulis complex in the North Atlantic region

Genetic relationships among Mytilus populations throughout the North Atlantic region, including the Mediterranean and the Baltic Sea, were studied using enzyme electrophoresis. Three distinct groups of populations, each of a remarkably wide distribution, can be recognised on the basis of their multilocus allelic composition: (1) M. galloprovincialis L. of the Mediterranean and western Europe; (2) a genetically distinct form of M. edulis Lmk. from both the Baltic Sea and some localities in the Canadian Maritime Provinces (here provisionally termed the trossulus type mussel); and (3) the traditional Atlantic M. edulis populations of northwestern European coasts and most of eastern North America. These groups are regarded as representing three relatively old evolutionary lineages, which all deserve separate and equal systematic status. The main part of the differentiation at most of the loci studied is accounted for by this major systematic pattern, but considerable geographical differentiation within each of the three principal groups was also detected. At single loci, different electromorphs were found to prevail in disjunct populations of M. galloprovincialis (Mediterranean/Britain) and of the trossulus-type mussel (Baltic/eastern Canada). Within the Atlantic M. edulis, a major part of the differentiation is transoceanic. At one locus (Ap), geographic differentiation appeared to be relatively independent of the systematic boundaries; the possible role of interlineage hybridisation in contact areas in regulating the pattern of geographical variation is discussed.     

Patterns of mtDNA diversity in North Atlantic populations of the mussel Mytilus edulis

Patterns of (female) mitochondrial DNA diversity were investigated in the blue mussel Mytilus edulis. Mytilus edulis is a ubiquitous member of contemporary North Atlantic hard-substrate communities and well represented in studies of this region. Mytilus edulis was surveyed in North America and Europe, as well as mid-Atlantic sites in Greenland, Iceland, and the Faroe Islands. Mitochondrial DNA sequences revealed considerable population structure but no monophyly of haplotypes between any major regions. Coalescent analyses suggest that migration across the Atlantic Ocean has prominently been from North American source populations and that Greenland was colonized recently and exclusively from North America. In North America, there was support for two regional groups along the North American coastline. Surprisingly, we also found evidence of recombination between some Mytilus edulis and Mytilus galloprovincialis female mtDNA sequences, particularly in northern Europe. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. An erratum to this article can be found at     

Genetic differentiation in Hediste diversicolor (Polychaeta: Nereididae) for the North Sea and the Baltic Sea

Genetic differentiation between North Sea and Baltic Sea Hediste diversicolor (O.F. Müller, 1776) (Polychaeta: Nereididae) populations was studied by allozyme electrophoresis on starch gel. Thirteen loci were analyzed in eight populations. The level of genetic variation was very low (mean H _o = 0.000 to 0.015). Differentiation between H. diversicolor populations is quite high (F _ST = 0.892) and reflected by three enzyme loci (MDH-I*, MDH-II*, IDH-I*). The reduced gene flow (N _m<1) may be explained by the limited dispersal capacity of the species. Regardless of whether found in the North Sea or Baltic Sea, there appear to be two different genetic types which are parapatric or sympatric in some places. The two types hybridize at three localities, but no signs of hybridization have been found at one (Tallinn). Received: 26 June 1997 / Accepted: 10 September 1997     

Ecological Costs of Livestock Grazing in Western North America

Livestock grazing is the most widespread land management practice in western North America. Seventy percent of the western United States is grazed, including wilderness areas, wildlife refuges, national forests, and even some national parks. The ecological costs of this nearly ubiquitous form of land use can be dramatic. Examples of such costs include loss of biodiversity; lowering of population densities for a wide variety of taxa; disruption of ecosystem functions, including nutrient cycling and succession; change in community organization; and change in the physical characteristics of both terrestrial and aquatic habitats. Because livestock congregate in riparian ecosystems, which are among the biologically richest habitats in arid and semiarid regions, the ecological costs of grazing are magnified in these sites. Range science has traditionally been laden with economic assumptions favoring resource use. Conservation biologists are encouraged to contribute to the ongoing social and scientific dialogue on grazing issues.     

Modelling potential habitat for cougars in midwestern North America

Cougars (Puma concolor) are of considerable interest to wildlife biologists and the general public in midwestern North America, yet no researchers have modelled potential habitat in the region. We created a model of potential cougar habitat in 9 midwestern states using geospatial data, expert-opinion surveys, the analytical hierarchy process, and a GIS. About 8% of the study region contained highly favorable habitat (with favorability scores ≥75%) for cougars; the states of Arkansas (19%) and Missouri (16%) contained the highest proportions of potentially favorable habitat. We identified 6 large (≥2500 km² in size), contiguous areas of highly favorable habitat for cougars. Model testing procedures indicated a valid model when compared to an independent set of cougar locations, a null dataset, and similar studies. Our model is useful as a planning tool to proactively address future human-cougar conflicts should cougars re-colonize the Midwest via subadult dispersal.     

Electrophoretic and biometric variability in the abyssal grenadier Coryphaenoides armatus of the western North Atlantic,eastern South Pacific and eastern North Pacific Oceans

Specimens of the abyssal grenadier Coryphaenoides armatus (Hector, 1875), from the western North Atlantic and eastern North Pacific Oceans were compared electrophoretically at 27 presumptive gene loci. At 6 of the 7 polymorphic loci there were only minor differences in allelic frequencies but a nearly fixed difference was found at one locus, phosphogluconate dehydrogenase. Eastern North Pacific grenadiers typically have a narrower interorbital space, a shorter dorsal interspace, more soft rays in the 1st dorsal fin (9–10 versus 8–9) and more pelvic fin rays (21–23 versus 18–21) than grenadiers from the western North Atlantic (as well as grenadiers from the eastern South Pacific, which were included in the biometric analysis). There is an apparent disjunction in the distribution of C. armatus in the eastern Pacific at the Gulf of Panamá which coincides with the change of morphology. It is suggested that North Pacific grenadiers comprise a subspecies, C. armatus variabilis Günther, 1878, which is morphologically distinct from the subspecies C. armatus armatus (Hector, 1875) of the other areas.     

Predicting Invasions of Woody Plants Introduced into North America

Plant species continue to be introduced in North America for various purposes. If the trend continues, it is probable that some will escape cultivation and become invasive in native ecosystems. We present a retrospective analysis of several structural, life history, and biogeographical attributes of woody plants introduced in North America to determine which traits characterize species that have and have not invaded. Predictive models derived from discriminant analysis correctly classified 86.2% of the species in cross-validation, whereas those derived from classification and regression trees classified 76% correctly. From these models we created a hierarchical predictive tree that allows the user to divide species into three categories: admit (low risk of invasiveness), deny admission (high risk of invasiveness), or delay admission for further analyses and/or monitor intensively (risk cannot adequately be assessed based on only the included attributes). We recommend that species that are highly invasive elsewhere not be allowed into the U.S. and that a more conservative introduction policy using a hierarchical predictive method be employed.     

Genetic differentiation in the brackish-water ostracod Cyprideis torosa

An electrophoretic survey of allozyme variation suggests that two separate populations of Cyprideis torosa (Jones) can be distinguished, one from the coast of England and the Netherlands, the other from the south coasts of the Baltic Sea. Gene flow between these two populations is limited and asymmetrical and seems to be more intense westwards than eastwards. On the other hand, particular local populations within the range of either population, although inhabiting geographically isolated estuaries, lagoons etc., are genetically similar to each other to such a degree that they should be treated as sets of subpopulations rather than as separate units. It is possible that in C. torosa a similar population structure occurs everywhere along the main migration routes of aquatic birds, which have clearly participated in the passive dispersal of this ostracod. Clines are observed in the frequencies of some alleles. Phosphogypsum pollution may be the cause of deviation from Hardy-Weinberg equilibrium in a local population.     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

Responses of Mytilus edulis on exposure to the water-accommodated fraction of North Sea oil

Individuals of Mytilus edulis L., collected from the Erme estuary (S.W. England) in 1978, were exposed to low concentrations (7 to 68 g l^-1) of the water-accommodated fraction (WAF) of North Sea crude oil. The pattern of accumulation of petroleum hydrocarbons in the body tissues was affected by the presence of algal food cells, the period of exposure, the hydrocarbon concentration in seawater, the type of body tissue and the nature of the hydrocarbon. Many physiological responses (e.g. rates of oxygen consumption, feeding, excretion, and scope for growth), cellular responses (e.g. lysosomal latency and digestive cell size) and biochemical responses (e.g. specific activities of several enzymes) were significantly altered by short-term (4 wk) and/or long-term (5 mo) exposure to WAF. Stress indices such as scope for growth and lysosomal latency were negatively correlated with tissue aromatic hydrocarbons.