The distribution of Melanoplus femurrubrum: fear and freezing in Connecticut

A central challenge for those studying organism distributions is to identify the relevant scale at which a distribution is established and the relevant mechanism driving the maintenance of the pattern. Melanoplus femurrubrum grasshoppers in NE Connecticut are missing from grass dominated fields. I examined two hypotheses about why the grasshopper is missing from these types of fields in this region. The first hypothesis was that behavioural responses to predation and competition prevent establishment by limiting above ground stages of the grasshopper. The second hypothesis was that soil temperature and grass habitat characteristics influence oviposition behaviour and limit egg development. Testing among these different hypotheses (trophic vs abiotic), data from experiments and field observations indicated that while the within field distribution of M. femurrubrum can be sensitive to the effects of trophic interactions during the above ground phase of the life-cycle, the among field distribution is limited by female oviposition choice, limited heating degree-days for egg development, and poor resource conditions for the first instar grasshoppers. These results highlight the need to examine behaviour and community interactions across habitats and life stages in studies of distribution patterns.     

Maternal environment and variation in ovariole number among populations of Melanoplus femurrubrum and M. scudderi scudderi

Ovariole number in univoltine populations of the grasshoppers Melanoplus femurrubrum (De Geer) and M. scudderi scudderi (Uhler) (Orthoptera: Acrididae) from Virginia was studied. Ovariole number in M. femurrubrum was found to vary among three geographic populations, and between years. Ovariole number did not vary between years in one population of M. s. scudderi. Ovariole number in M. femurrubrum was positively correlated with the mean length of the growing season at each site and precipitation during August-September of the previous year (adult season of the maternal generation). In the laboratory, nymphal rearing temperatures did not affect ovariole number in M. femurrubrum, but it was found to be weakly correlated with egg size.

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Résumé Le nombre d'ovarioles des criquets détermine le nombre maximum d'oeufs par ponte. Si la variabilité du nombre d'ovarioles des criquets a souvent été signalée, l'origine de ces variations n'a été examinée que pour quelques espèces migratrices, et cela, au laboratoire.Le nombre d'ovarioles des populations monovoltines de M. femurrubrum a changé suivant les populations et suivant les années. Une étude par régressions multiples a montré que le nombre d'ovarioles est lié à des caractéristiques écologiques de la génération maternelle: durée de la saison de développement, indiquée par la somme des températures au-dessus de 10°C, et précipitations en août et septembre. La durée de la saison de développement détermine le temps disponible pour la reproduction: les populations, aux saisons de développement les plus longues, ont plus de temps pour former une oothèque, et ainsi disposent des moyens métaboliques pour avoir plus d'ovarioles. Août-septembre est l'époque où les criquets deviennent adultes, commencent à s'alimenter avant la ponte, et pondent la majorité de leurs oothèques. On estime que l'effet des précipitations est indirect, par son action sur la quantité et la qualité des plantes-hôtes disponibles. Le nombre d'ovarioles n'est pas lié à la taille de M. femurrubrum ou de M. scudderi scudderi.Les effectifs de M. s. scudderi étaient insuffisants pour permettre de telles analyses; cependant, nous n'avons observé dans une station aucune modification du nombre d'ovarioles au cours des années.

     

Genetic divergence of insular populations of deer mice

Electrophoretic variants at 28 genetic loci were analyzed in subspecies of Peromyscus maniculatus endemic to the Channel Islands off the California coast. The genetic variability within insular populations was calculated. These deer mice have relatively high levels of polymorphism for insular populations. The mean heterozygosity per individual varies in the populations, being much higher on one of the islands than the others. Nei's measure of genetic distance between groups compared on the basis of electrophoretic variants was used. His estimate of time of divergence of these groups, based on genetic distance, is applicable particularly to closely related groups. The length of time each island population has been isolated from the others was calculated and found to be consistent with the periods of isolation estimated on the basis of geological data.     

Feeding preferences of Melanoplus femurrubrum grasshoppers on native and exotic grasses: behavioral and molecular approaches

Generalist insect herbivores, such as grasshoppers, may either avoid feeding on exotic plants, potentially enabling these plants to become invasive in the introduced range, or insects may incorporate exotic plants into their diet, contributing to the biotic resistance of native communities and potentially preventing plant invasions. Accurate determination of insect diet preferences with regard to native and exotic plants can be challenging, but this information is critical for understanding the interaction between native herbivores and exotic plants, and ultimately the mechanisms underlying plant invasions. To address this, we combined behavioral and molecular approaches to accurately compare food consumption of the polyphagous red‐legged grasshopper, Melanoplus femurrubrum (De Geer) (Orthoptera: Acrididae), on native [Andropogon gerardii Vitman and Bouteloua curtipendula (Michx.) Torr.] and exotic, potentially invasive grasses [Miscanthus sinensis Andersson and Bothriochloa ischaemum (L.) Keng] (all Poaceae). We found that M. femurrubrum grasshoppers demonstrated strong feeding preferences toward exotic grasses in experiments with intact plants under both field and greenhouse conditions, but they showed no preference in experiments with clipped leaves. Additionally, we sampled the gut contents of M. femurrubrum collected in the field and identified the ingested plant species based on DNA sequences for the non‐coding region of the chloroplast trnL (UAA) gene. We found that exotic plants were prevalent in the gut contents of grasshoppers collected at study sites in Ohio and Maryland, USA. These results suggest that the generalist herbivore M. femurrubrum does not avoid feeding on exotic grasses with which they do not share coevolutionary history. In addition, by demonstrating greater food consumption of exotic plants, these grasshoppers potentially provide biotic resistance should these grasses escape cultivation and become invasive in the introduced range.     

Genetic and behavioural divergence among desert spring amphipod populations

1. Random amplification of polymorphic DNA (RAPD) analyses and behavioural trials were conducted on eleven Hyalella azteca populations and one population of H. montezuma from lakes and springs across northern Arizona, U.S.A. Genetic distances among populations were calculated based on fifty-five RAPD markers.
2. A cluster analysis indicated two genetic lineages among the Hyalella populations. One group consisted of populations inhabiting lakes with submerged vegetation, while a second group consisted of populations that clung to roots of emergent vegetation. Behaviour also diverged between these two groups in laboratory trials: amphipods from the submerged vegetation lineage spent significantly more time swimming than did those from the emergent vegetation lineage.
3. Based on patterns in genetic distance data, it is suggested that a xeric landscape may promote diversity among passively dispersed invertebrates.     

Genetic structure and divergence of tench Tinca tinca European populations

The tench Tinca tinca is a freshwater species with human-mediated translocations, aquaculture interest and limited information on its genetic structure. mtDNA sequencing analysis of control region and two genes in 50 individuals from five European populations identified two phylogroups, with greater variability than that reported until now, and a hybridization zone in the Danube River region. Restriction analyses of additional samples reveal the complicated genetic structure characteristics of tench's wild and translocated populations, supporting future breeding practices.     

Genetic divergence without spatial isolation in polecat Mustela putorius populations

Understanding how genetic divergence could exist without spatial isolation is a fundamental issue in biology. Although carnivores have previously been considered as having a weak genetic variability, polecats Mustela putorius from eight distinct populations exhibited both a strong polymorphism (17.5–22.5%) and a substantial allele effective number reaching N_e=1.12. Heterozygosity ranging from H_o=0.031–0.063 significantly differed among populations, while the mean F_IS averaging 0.388 stressed a real deficiency of heterozygotes. Observed heterozygosity levels among populations did not correlate with any habitat types but were clearly associated with habitat diversity index. The habitat structure in polecat home range corresponded to habitat mosaic structure in which discrete habitat types alternated causing multifactorial constraints that may favour heterozygosity. Allozymic frequencies within populations did not vary with dominant habitat. But in the Tyrosinase‐1, the rare homozygote BB, resulting in a ‘dark’ phenotype, was found much more in deciduous woods than the homozygote AA showing the ‘typical’ pattern. Thus, the genetic basis for a character differentiation was here evidenced in a remarkable situation without spatial isolation. Further, the very low proportion of heterozygotes for this locus suggests a disruptive effect and supports the prediction of intermediate phenotypes being at a disadvantage. This heterozygote deficit may also result from an assortative mating intra phenotype (homogamy). The divergence in polecat phenotypes showed that genetic differentiation can be induced by subtle variations in environment, a situation that is likely to be frequent in most natural populations, and emphasized the adaptive nature of habitat preference.     

Consumption rates and the evolution of diet-induced plasticity in the head morphology of Melanoplus femurrubrum (Orthoptera: Acrididae)

Summary Phenotypic plasticity may be an ecologically important evolutionary response to natural selection in multiple environments. I have determined the effect of diet-induced developmental plasticity in the head size of grasshoppers (Melanoplus femurrubrum) onfeeding performance on two types of plants. Full-sib families were divided and raised on either red clover, Trifolium repens, or rye grass, Lolium perenne. In three different stages of ontogeny, grasshoppers raised on rye grass had significantly larger heads, relative to body size, than full-sibs raised on clover. A principal components analysis indicated that two to five relative head size characters covaried as a block in their plastic response to the feeding environment. Regressions of adjusted consumption rates (mg/sec) against relative head size revealed that larger head sizes, induced by the rye grass diet, enhanced consumption rates of rye grass, but not clover. Unexpectedly, a similar positive association was observed between head size and consumption rate for grasshoppers raised on clover when they were feeding on clover. These results support the inference that grasshoppers exhibit adaptive phenotypic plasticity. However, the unexpected influence of head size on consumption rates of clover indicates that the functional relationship between head morphology and feeding performance is complex and that variation in this relationship among plant environments is not sufficient to explain the evolution of diet-induced phenotypic plasticity.     

Genetic divergence among geographical populations of the migratory locust in China

The random amplified polymorphic DNA (RAPD) technique was used to examine genetic divergence and interrelations of 11 geographical populations of the migratory locust in China, and the role of spatial separation in the population differentiations. AMOVA analysis of genetic variations in all the populations indicated greater within- (79.55%) than among-population variability (20.45%), and that there were significant differentiations among the populations; 11 populations were divided into four regional groups, with significantly greater variability within (82.99%) than among the groups (17.01%), and there existed apparent regional differentiations. Paired comparisons showed significantly greater variability within-than between-groups, indicating significant differentiations between populations of different regional groups. Of all the pairwise comparisons, Hainan and Tibetan groups displayed the greatest differentiation, with the difference between the two groups being seven folds of that between populations within the groups; the least differentiations were exhibited between the groups of Hainan, Xinjiang, and Inner Mongolia, with the differences between groups being only half of the differences between populations within the groups. Mantel tests of the genetic and spatial distances showed that the two matrices were significantly correlated (p<0.01), indicating that the geographical isolation played an important role in the differentiations of the geographical populations of the migratory locusts. Cluster analysis divided all populations into four major groups: Xinjiang and Inner Mongolia group, the Great Plains of North China (the Yellow River and Huai River Plains) group, Hainan group, and Tibet group. Principal component analysis (PCA) supported the division of populations based on the cluster analysis. However, analysis of individuals clustered the locusts into five populations: Xinjiang and Inner Mongolia, Hami in Xinjiang, the Great Plains of North China, Hainan, and Tibet. The locust populations in eastern China displayed apparently continous and gradient variations; as such authors consider that there were no necessity and valid reasons for further division of subspecies. The subspecific status for the main geographical populations of the migratory locusts in China was discussed.     

Genetic divergence among geographical populations of the migratory locust in China

The migratory locust, Locusta migratoria L., which is distributed widely in the East Hemisphere, has long been regarded as the most important agricultural pest and a model insect. Its distribution ranges from thetropic and the subtropical zone, to the temperate zone, and to the cold-temperate zone. Because of the wide geographical distribution and adaptations, many geo- graphical populations of the migratory locust dis- played apparent variation in morphology, life history, physiology, and ot…     

Genetic variability and divergence between populations and species of Nesticus cave spiders

Genetic variability and divergence at 21 enzyme loci were studied in and between Italian populations of the cave spiders Nesticus eremita (13 populations), N. menozzii and N. sbordonii (one population each). The three species differ with respect to the degree of specialization to cave life, dispersion ability, isolation of populations, abundancy, extent of the distribution area, and range from the troglophilic and widespread N. eremita to the troglobitic N. sbordonii, endemic to a single cave in the Central Appennines.Heterozygosity ranges from 0.05 to 0.15 in N. eremita populations and appears to be largely controlled by the occurrence and the extent of gene flow among populations. The relatively low polymorphism levels of N. menozzii (H=0.081) and N. sbordonii (H=0.106) are also associated with reduced gene flow and small population sizes.Genetic distances between N. eremita populations vary considerably and are strictly related to the geographical distances involved, again indicating a major role of gene flow in determining the patterns of genetic differentiation between populations. This view is strongly supported by the results of a principal component analysis applied to the gene-frequency data. Estimates of genetic divergence between species suggest that the major cladogenetic events leading to complete separation of these three Nesticus species occurred in the Middle-late Pliocene.     

Genetic divergence between Melipona quadrifasciata Lepeletier (Hymenoptera,Apidae) populations

Melipona quadrifasciata is a stingless bee widely found throughout the Brazilian territory, with two recognized subspecies, M. quadrifasciata anthidioides, that exhibits interrupted metasomal stripes, and M. quadrifasciata quadrifasciata, with continuous metasomal stripes. This study aimed to estimate the genetic variability of these subspecies. For this purpose, 127 colonies from 15 Brazilian localities were analyzed, using nine species-specific microsatellite primers. At these loci, the number of alleles ranged from three to 15 (mean: 7.2), and the observed heterozygosity (H_o) ranged from 0.03–0.21, while the expected heterozygosity (H_e) ranged from 0.23–0.47. The genetic distances among populations ranged from 0.03–0.45. The F_ST multilocus value (0.23) indicated that the populations sampled were structured, and the clustering analysis showed the formation of two subgroups and two more distant populations. The first group contained the subspecies M. quadrifasciata quadrifasciata, and the other, the subspecies M. quadrifasciata anthidioides and the two M. quadrifasciata populations with continuous metasomal stripes from northern Minas Gerais. These results confirmed that the yellow metasomal stripes alone are not a good means for correctly identifying the different subspecies of M. quadrifasciata.     

Genetic divergence of climatically marginal populations of Vicia pisiformis on the Scandinavian Peninsula

Vicia pisiformis L. is a perennial leguminous plant with a main distribution in broadleaved forest-steppes of eastern Europe. The species is classified as endangered (EN) according to the IUCN red-lists in both Norway and Sweden, due to severe fragmentation, small population sizes and continuing population decline. The populations on the Scandinavian Peninsula constitute the northern limit of the species distribution and are mostly restricted to warm stony slopes with predominantly southern aspects. In this study we used the AFLP method, which is a high-resolution genetic fingerprint method. Samples were collected from 22 Scandinavian populations. The overall genetic structure was analysed in an AMOVA, in a Mantel test and through constrained correspondence analysis (CCA). The ordination scores representing non-geographic genetic divergence were extracted from the CCA and analysed in a linear model using habitat variables and population size as explanatory variables. We found (i) a strong geographic structure, (ii) significant genetic divergence between populations, (iii) that this genetic divergence remained significant even after removing the effect of geography in a partial CCA and (iv) that the remaining non-geographic part of genetic divergence (distance from the ordination centre) was associated with aspect, populations with a northern aspect were more genetically divergent. Aspect explains more variation than population size and is the only variable retained in the minimal adequate model. We suggest that local adaptation has caused this divergence from an expected geographical pattern of genetic variation. This explanation is further supported by the association between aspect and specific AFLP fragments.     

Genetic divergence and isolation by thermal environment in geothermal populations of an aquatic invertebrate

Temperature is one of the most influential forces of natural selection impacting all biological levels. In the face of increasing global temperatures, studies over small geographic scales allowing investigations on the effects of gene flow are of great value for understanding thermal adaptation. Here, we investigated genetic population structure in the freshwater gastropod Radix balthica originating from contrasting thermal habitats in three areas of geothermal activity in Iceland. Snails from 32 sites were genotyped at 208 AFLP loci. Five AFLPs were identified as putatively under divergent selection in Lake Mývatn, a geothermal lake with an almost 20 °C difference in mean temperature across a distance of a few kilometres. In four of these loci, variation across all study populations was correlated with temperature. We found significant population structure in neutral markers both within and between the areas. Cluster analysis using neutral markers classified the sites mainly by geography, whereas analyses using markers under selection differentiated the sites based on temperature. Isolation by distance was stronger in the neutral than in the outlier loci. Pairwise differences based on outlier F_ST were significantly correlated with temperature at different spatial scales, even after correcting for geographic distance or neutral pairwise F_ST differences. In general, genetic variation decreased with increasing environmental temperature, possibly suggesting that natural selection had reduced the genetic diversity in the warm origin sites. Our results emphasize the influence of environmental temperature on the genetic structure of populations and suggest local thermal adaptation in these geothermal habitats.     

Genotype–environment interaction and the ontogeny of diet-induced phenotypic plasticity in size and shape of Melanoplus femurrubrum (Orthoptera: Acrididae)

The developmental origin of phenotypic plasticity in morphological shape can be attributed to environment-specific changes in growth of overall body size, localized growth of a morphological structure or a combination of both. I monitored morphological development in the first four nymphal instars of grasshoppers (Melanoplus femurrubrum) raised on two different plant diets to determine the ontogenetic origins of diet-induced phenotypic plasticity and to quantify genetic variation for phenotypic plasticity. I measured diet-induced phenotypic plasticity in body size (tibia length), head size (articular width and mandible depth) and head shape (residual articular width and residual mandible depth) for grasshoppers from 37 full-sib families raised on either a hard plant diet (Lolium perenne) or a soft plant diet (Trifolium repens). By the second to third nymphal instar, grasshoppers raised on a hard plant diet had significantly smaller mean tibia length and greater mean residual articular width (distance between mandibles adjusted for body size) compared with full-sibs raised on a soft plant diet. However, there was no significant phenotypic plasticity in mean unadjusted articular width and mandible depth, and in mean residual mandible depth. At the population level, development of diet-induced phenotypic plasticity in grasshopper head shape is mediated by plastic changes in allocation to tissue growth that maintain growth of head size on hard, low-nutrient diets while reducing growth of body size. Within the population, there was substantial variation in the plasticity of growth trajectories since different full-sib families developed phenotypic plasticity of residual articular width through different combinations of head and body size growth. Genetic variation for diet-induced phenotypic plasticity of residual articular width, residual mandible depth and tibia length, as estimated by genotype–environment interaction, exhibited significant fluctuation through ontogeny (repeated measures MANOVA , family × plant × instar, P < 0.01). For example, there was significant genetic variation for phenotypic plasticity of residual articular width in the third nymphal instar, but not earlier or later in ontogeny. The observed patterns of genetic variation are discussed with reference to short-term constraints and the evolution of phenotypic plasticity.     

Genetic divergence does not predict change in ornament expression among populations of stalk-eyed flies

Stalk-eyed flies (Diptera: Diopsidae) possess eyes at the ends of elongated peduncles, and exhibit dramatic variation in eye span, relative to body length, among species. In some sexually dimorphic species, evidence indicates that eye span is under both intra- and intersexual selection. Theory predicts that isolated populations should evolve differences in sexually selected traits due to drift. To determine if eye span changes as a function of divergence time, 1370 flies from 10 populations of the sexually dimorphic species, Cyrtodiopsis dalmanni and Cyrtodiopsis whitei, and one population of the sexually monomorphic congener, Cyrtodiopsis quinqueguttata, were collected from Southeast Asia and measured. Genetic differentiation was used to assess divergence time by comparing mitochondrial (cytochrome oxidase II and 16S ribosomal RNA gene fragments) and nuclear (wingless gene fragment) DNA sequences for c. five individuals per population. Phylogenetic analyses indicate that most populations cluster as monophyletic units with up to 9% nucleotide substitutions between populations within a species. Analyses of molecular variance suggest a high degree of genetic structure within and among the populations; > 97% of the genetic variance occurs between populations and species while < 3% is distributed within populations, indicating that most populations have been isolated for thousands of years. Nevertheless, significant change in the allometric slope of male eye span on body length was detected for only one population of either dimorphic species. These results are not consistent with genetic drift. Rather, relative eye span appears to be under net stabilizing selection in most populations of stalk-eyed flies. Given that one population exhibited dramatic evolutionary change, selection, rather than genetic variation, appears to constrain eye span evolution.     

The genetic divergence of three populations

This paper considers the effect of the genetic divergence on the genetic composition of three samples drawn from three populations at some time after the populations had split. It generalizes the two-sample case studied earlier by Watterson (1985a). Under the assumptions that (i) mating is at random, (ii) the genes at a locus can be any of infinitely many alleles and all mutants are assumed to be new alleles, and (iii) no selective differences exist, we find the probability distribution of the sample gene configurations. From this distribution the single-sample allelic distribution after one-step and two-step bottlenecks and the allelic distribution in the two-sample case can be obtained as marginal distributions. Some numerical results on the number of alleles in common in the three samples are compared with those obtained by Watterson's simulation method; the agreement is excellent. Also, the probability that the three samples are monomorphic for the same allele is found, and numerical examples are given.     

The genetic divergence of two populations

When a population splits in two and the resulting populations evolve independently, genetic divergence may occur due to mutation and genetic drift. This paper considers the influence these factors have on the genetic composition of two samples drawn from the respective populations at some time after the split. Analytical formulas and numerical examples are given of (i) the probability distribution and moments of the allele frequencies, (ii) the number of alleles in common between the samples, and (iii) the probability that the samples are monomorphic.     

Genetic divergence in Greek populations of the genus Leuciscus and its evolutionary and biogeographical implications

In 23 populations of Greek Leuciscus ( Squalius ), the percentage of polymorphic allozyme loci ranged between 0·034 and 0·379 (P=0·19) and expected heterozygosity was 0·011–0·166 (H_e=0·067). Current taxonomy is confusing and does not correspond to genetic data that support the presence in Greece of at least seven different species: L. cephalus , L. peloponnensis , L. prespensis , L. moreoticus , L. borysthenicus , L. keadicus and Leuciscus sp. from Euboea Island. The maximum Nei modified genetic distance was found among L. keadicus and the rest of subgenus Squalius populations D * Nei=0·446–0·705). Accepting the molecular clock hypothesis, speciation for the genus Leuciscus in Greece must have occurred during the Cenozoic period (between the Middle Miocene and the Later Pliocene). The two main biogeographical events causing speciation on mainland Greece were the Uplift of the Pindic cordillera and the isolation of the southern part of Peloponnesus. The faunistic composition of the lakes studied, in which new taxa are reconsidered, suggests the same faunistic origin in the Early Pliocene for Lakes Prespa and Stymphalia and a younger one in the Late Pliocene for Lake Trichonis. Euboea Island was not a zoogeographical unit during the Cenozoic. The isolation of all the freshwater fish fauna of Euboea has occurred since the Pliocene. The biogeographical model proposed here differs from classical hypotheses in considering of lesser importance the dispersion of L. cephalus on the Greek mainland during the Neogene and Quaternary.