Genetic Status and Fluctuating Asymmetry in an Endangered Population of the Moth Dysauxes ancilla L. (Lepidoptera: Ctenuchidae)

Genetic status and fluctuating asymmetry (FA) were assessed in a small, isolated and endangered population of the moth Dysauxes ancilla in Sweden. A sample from the German population, within the continuous breeding area of the species, was used for comparison. The levels of FA were significantly higher in the Swedish population, indicating a reduced ability to withstand developmental stress. Two polymorphic loci showed significantly higher variation in the Swedish population, indicating that there are no serious effects on genetic factors. Therefore, it is suggested that the increased level of FA in the Swedish population is due to the stress of living in an ecologically marginal habitat. The Swedish population is a northern outpost separated from the continuous distribution area of species and environmental stress caused by variable and extreme abiotic factors, for example climatic conditions, could explain a higher FA. However, it is still an open question if a higher FA from environmental stress also constitutes an increased extinction risk.     

Fluctuating Asymmetry and Male Mating Success in a Sphragis-Bearing Butterfly Luehdorfia japonica (Lepidoptera: Papilionidae)

The relationship between fluctuating asymmetry (FA) and mating success was studied within males of the sphragis-baring butterfly Luehdorfia japonica, which were collected at various periods during their mating season. FA was measured on the forewing and hindwing radius lengths of male butterflies. Mating frequency of males was estimated by assessing the degree of scale loss from their claspers. Males consume scales and use them to form sphragis on the female abdomen during copulation, sealing the ostium bursa for life. Age of males was scored as wing age 0 to 4 according to the wearing of the wing. FA was negatively correlated with mating frequency but positively correlated with wing age, and average FA decreased with mating season. As females have little chance to express mate choice, it is likely that FA is an indicator of male viability: symmetrical males live longer and/or fly more actively, resulting in a higher lifetime mating success compared to asymmetrical males.     

Experimental Extinction of Garlic Mustard (Alliaria petiolata) Populations: Implications for Weed Science and Conservation Biology

An assumption of weed science and conservation biology is that small populations are more vulnerable to elimination and extinction than large populations. We tested this with the invasive biennial garlic mustard (Alliaria petiolata). We compared 61 experimental populations from which every flowering plant was removed for 4 years, with 56 control populations. Whereas the majority of the control populations continued to expand in size over the 4 years, experimental populations showed a strong experimental effect, remaining stable in size, declining in size, or going extinct. Small populations were far more vulnerable to extinction than large populations: 43% of small experimental populations (initially fewer than 10 individuals) went extinct, but only 7% of large populations (initially more than 50 individuals). However, some small experimental populations persisted, and in a few cases, larger experimental populations continued to expand even though every flowering individual had been removed. These results and a simple population model suggest the importance of buried seeds in allowing this species to persist despite attempts to eradicate it.     

Fluctuating asymmetry inMacaca fascicularis: A study of the etiology of developmental noise

Fluctuating asymmetry was determined for six cranial measurements in an age-diverse sample of 138 individuals ofMacaca fascicularis. These data were used to choose among four hypotheses concerning the etiology of developmental noise. The hypotheses considered are (1) that developmental noise represents asymmetry in the causal history of a developing organism's interaction with the environment, (2) that it represents stochasticity in the mechanics of growth and induction, (3) that it reflects variation in the initial conditions of a developmental process, and (4) that it represents the random accumulation of noise at a level below that of morphogenetic mechanism. These hypotheses were tested against predictions concerning the intraspecific patterning of fluctuating asymmetry against age and size and the covariation of asymmetry values. Only the predictions of the fourth hypothesis were confirmed by results of this study. These results provide evidence for the view that developmental noise, as reflected by fluctuating asymmetry, is an intrinsic property of developmental systems, and not merely produced by the complexity of the organism's interaction with the environment.     

Ontogenetic Patterning of Skeletal Fluctuating Asymmetry in Rhesus Macaques and Humans: Evolutionary and Developmental Implications

I address the question of how fluctuating asymmetry (FA)—the distribution of random deviations from bilateral symmetry—varies ontogenetically in the mammalian skeleton. This question is significant because of the light that such patterns can shed on the causes of variation in developmental stability in bone as well as other structures. Based on large ontogenetic skeletal series of Macaca mulatta and Homo sapiens, I report that the FA variances of skeletal metric traits increase ontogenetically. Coupled with the finding that FA variances also accumulate to greater magnitudes in slower growing mammals, this result is consistent with the hypotheses that FA in bone is primarily caused by (a) cumulative effects of asymmetrical mechanical factors, (b) accumulation of variation in the (local) regulation of growth, or (c) a tendency for bone morphology to drift through undirected remodeling. I discuss the implications of these optional explanations for primate evolution and bone development.     

Fluctuating asymmetry for specific bristle characters in Notch mutants of Drosophila melanogaster

Asymmetry has been used as a measure of developmental stability for bilaterally symmetrical organisms. Most studies have failed to partition the genetic and environmental contributions to the asymmetry phenotype due to the limitations of the systems used or the shortcomings in experimental design. The Notch mutants of Drosophila melanogaster were used to study the genetic contribution to asymmetry for six different bristle characters. Asymmetry response was character specific for the mutants examined. For N ^spl, N ^Co, N ^264–47, Ax ^71d, Ax ^9B2, Ax ^E2, l(1)N ^B and nd ² significant asymmetry responses, relative to wildtype Canton‐S, were observed for some characters. N ^60g11 and nd ¹ did not exhibit significant asymmetry for any of the characters examined. All of the mutants except N ^60g11 and nd ¹ showed thoracic bristle asymmetry. However, when asymmetry scores were pooled over the five bristle characters which individually exhibited fluctuating asymmetry, no significant differences were found between any genotypes. Therefore pooling asymmetry values across characters obscures the significant character specific asymmetry values observed. Thus caution is necessary when using the asymmetry phenotype of specific characters to draw organism wide conclusions about developmental stability. This revised version was published online in July 2006 with corrections to the Cover Date.     

Fluctuating Asymmetry in Body Traits Increases Predation Risks: Tawny Owl Selection Against Asymmetric Woodmice

During the last decade, the study of fluctuating asymmetry (FA) in relation to different fitness aspects has become a popular issue in evolutionary biology. There has been much recent debate in subtle departures from perfect symmetry in bilaterally paired morphological characters, and the extent to which such departure actually reflects aspects of individual quality and fitness. We used data from pellet collection and trapping sessions involving the trophic system Apodemus – Strix aluco, to test the hypothesis that asymmetric woodmice disproportionately fell prey to the tawny owl compared with “normal” woodmice. We found that woodmice preyed on by owls had significantly more asymmetric leg bones than survivors, particularly hind legs, those devoted to jumping. Thus asymmetry in locomotory traits apparently increased predation risks due probably to minor efficiency of asymmetric woodmice in evading predators or to their general low quality. These results suggest that FA affects fitness and consequently may be a good predictor of survival chances for woodmice, i.e. their quality; on the other hand, by removing asymmetric individuals, tawny owls can exert a stabilising selection on prey populations.Co-ordinating editor: M. Klaassen     

Conservation Biology of the Genus Alouatta

As assessed by the IUCN Mace–Lande system, seven (35%) of the 20 Alouatta species and subspecies with adequate data are classified as threatened, i.e., critically endangered, endangered, or vulnerable (Rylands et al., 1995). This percentage is much lower than the 75 to 100% threatened taxa for the other large-bodied genera: Ateles, Lagothrix, and Brachyteles. Only 5 of the 16 Neotropical genera have lower percentages of threatened taxa than that of Alouatta: Cebuella, Pithecia, Saguinus, Saimiri, and Cebus. The threatened howler taxa occupy small distributions in areas of forest fragmentation. In general, populations are most affected by major habitat disturbance, such as total deforestation and flooding from dam construction, and by human hunters. Facilitated by their ability to exploit folivorous diets in small home ranges, howlers can tolerate considerable habitat fragmentation but not the increased exposure to hunting that may accompany it. Howlers seem particularly vulnerable to yellow fever and bot fly parasitism. Although the former threat may decrease by increasing fragmentation of the habitat, other sorts of parasitism may increase in disturbed habitats. The low genetic variability of the Central American howlers suggests a resistance to inbreeding depression potentially experienced during population bottlenecks. Greater between-population variability may still exist. Although howlers are not readily bred in captivity, they respond well to translocation. Translocation has been successfully achieved for 4 howler species and is a viable option for introducing new genetic variability into population fragments and repopulating areas from which howlers are extinct. Their pattern of bisexual dispersal facilitates colonization of regenerating habitats, and in suitable, protected habitats they have shown the capacity for strong population recovery.     

Flight and Oviposition Behavior of the Adult Maritime Ringlet (Coenonympha nipisiquit McDunnough) Females in Response to Microhabitat

Maritime ringlet butterflies (Coenonympha nipisiquit McDunnough), an endangered species in Canada, inhabit salt marshes, which consist of microhabitat mosaics with varied larval survival rate. These microhabitats may influence the movement and reproductive behaviors of females, which in turn may affect population dynamics. I recorded behaviors and locations of females every minute with a GPS rover and calculated their move lengths and turning angles. Move lengths did not change in response to microhabitats, although turning angles became larger near bodies of water with sparse vegetation. Females spent a longer time in one location and oviposited more often where the principal larval host, Spartina patens (Aiton) Muhl., is abundant, regardless of larval survival rate. Older females tended to initiate flight more readily than younger females and spent more time flying and nectaring. Younger females were more fecund and spent a longer time at one location. Because young females tend to be less mobile and more fecund, the majority of oviposition should take place near eclosion sites. However, some eggs will be laid away from microhabitats favorable to larval survival when older females become mobile and move out of their natal microhabitats. Because it seems to have little potential to colonize new habitat on its own, monitoring population dynamics and habitat quality will be crucial for the persistence and recovery of this rare species.     

Fluctuating asymmetry and survival ability in the forest tent caterpillar mothMalacosoma disstria: implications for pest management

Fluctuating asymmetry of the first tarsal segment of the proleg of the forest tent caterpiller mothMalacosoma disstria Hbn. (Lepidoptera: Lasiocampidae) was significantly inversely related to survival ability in the lab. The monitoring of population levels of fluctuating asymmetry could have important implications in pest management of this and other species by providing an indication of the health of a population.     

Perfect is best: low leaf fluctuating asymmetry reduces herbivory by leaf miners

Fluctuating asymmetry (FA) represents small, random variation from symmetry and can be used as an indicator of plant susceptibility to herbivory. We investigated the effects of FA of two oak species, Quercus laevis and Q. geminata, and the responses of three herbivore guilds: leaf miners, gallers, and chewers. To examine differences in FA and herbivory between individuals, 40 leaves from each tree were collected, and FA indices were calculated. To examine differences in FA and herbivory within-individuals, we sampled pairs of mined and unmined leaves for asymmetry measurements. Differences in growth of leaf miners between leaf types were determined by tracing 50 mines of each species on symmetric leaves and asymmetric leaves. Asymmetric leaves contained significantly lower concentrations of tannins and higher concentrations of nitrogen than symmetric leaves for both plant species. Both frequency of asymmetric leaves on plants and levels of asymmetry positively influenced the abundance of Brachys, Stilbosis and other leaf miners, but no significant relationship between asymmetry and herbivory was observed for Acrocercops. Brachys and Stilbosis mines were smaller on asymmetric leaves, but differences in mine survivorship between symmetric and asymmetric leaves were observed only for Stilbosis mines. This study indicated that leaf miners might use leaf FA as a cue to plant quality, although differential survivorship among leaf types was not observed for all species studied. Reasons for the different results between guilds are discussed.     

Modification of developmental instability and fitness: Malathion-resistance in the Australian sheep blowfly,Lucilia cuprina

The evolution of resistance to malathion byLucilia cuprina initially results in an increase in fluctuating asymmetry. Resistant flies are at a selective disadvantage, relative to susceptibles, in the absence of the insecticide. A fitness/asymmetry modifier of diazinon-resistant phenotypes ameliorates these effects resulting in malathion-resistant phenotypes of relative fitness and asymmetry similar to susceptibles. For the nine genotypic combinations of the modifier and malathion-resistance alleles, developmental time increases linearly with increasing asymmetry. Percentage egg hatch decreases linearly with increasing asymmetry. The initially disruptive effect of the malathion-resistant allele was partially dominant, the effect of the modifier dominant. The results are discussed in terms of developmental perturbation, asymmetry estimation and relative fitness to consider whether it is adequate to use changes in fluctuating asymmetry alone as measures of developmental instability. It is suggested that in some circumstances antisymmetry may indicate developmental instability and that the diazinon/malathion-resistance systems inL. cuprina may allow the relative importance of genetical and/or environmental developmental perturbations to be ascertained.     

Conservation of Drill Populations in Bakossiland,Cameroon: Historical Trends and Current Status

Very little information is available on the current status of drill populations in Cameroon. We report on drill group sizes and status in Bakossiland, a mountainous area spanning 2000 km² in the Littoral and South West Provinces of southwestern Cameroon. Between 1970 and 2002 direct visual counts of drill groups (n = 105) yielded group size estimates ranging from 5 to 400 (mean ± S.E.M.; 93.1 ± 8.4). We encountered solitary adult male drills on 8 occasions. Groups were at all elevations (150–2000 m) in 5 habitat types: lowland, premontane, submontane and montane forests and montane savannah at 2000 m). Group sizes did not vary with respect to elevation, habitat type or season (wet and dry mo). However, over the past decade drills have been virtually hunted out of the Mwenzekong Mountains (Banyang Mbo Wildlife Sanctuary), and they are greatly reduced in the southern Bakossi forests of Mwendolengo, Edib Hills and Mungo River. The species became extinct in the Loum Forest Reserve in the late 1970s, and until recently was thought to have become extinct on Mount Mwanenguba. Since 1994 on Mount Kupe, the drill population has begun to recover, largely due to protection afforded by the Bakossi traditional chiefs. Traditional powers and values are still influential in the region. A new national park—Bakossi Mountains National Park— and associated Protected Areas are currently under gazettment. We discuss the effectiveness of conservation strategies in relation to the survival of drills in the area.     

The Influence of Weapon Asymmetry on Male-Male Competition Success in a Sexually Dimorphic Insect,the African King Cricket Libanasidus vittatus (Orthoptera: Anostostomatidae)

Journal of Insect Behavior -     

Biology of Lilioceris spp. (Coleoptera: Chrysomelidae) and their parasitoids in Europe

The biology and parasitoid complex of the lily leaf beetle (LLB), Lilioceris lilii Scopoli, and two congeneric species were investigated in Europe, as part of a biological control program against the LLB in North America. Eggs, larvae, and adults of L. lilii were collected in several countries in Europe, on both cultivated and wild Lilium spp., and reared in the laboratory and under natural conditions. Parasitoids were obtained and their biologies were studied. Similar investigations were made in Switzerland on two closely related species Lilioceris tibialis (Villa) found on wild Lilium spp., and Lilioceris merdigera (L.) on several other Liliaceae. The three species are strictly univoltine. Adults overwinter and lay eggs on leaves in early spring. The three beetle species have four instars, which were characterized by their head capsule width. Pupation occurs in a cocoon in the soil. Adults emerge in late summer and start feeding before reaching overwintering sites. Egg and larval parasitoids were obtained. Eggs of L. lilii and L. merdigera were parasitized by the mymarid Anaphes sp., a multivoltine species that needs alternate hosts for overwintering. Larvae were heavily attacked by several parasitoids, among which the most abundant were three ichneumonids, Lemophagus pulcher (Szepligeti), L. errabundus (Gravenhorst), and Diaparsis jucunda (Holmgren), and the eulophid Tetrastichus setifer Thomson. All four parasitoid species were found in the three beetles and in most European regions, but strong variations were observed in their relative abundance among hosts and geographic regions. Three of the four main larval parasitoids are strictly univoltine, whereas L. pulcher has a partial second generation. Lemophagus spp. are frequently parasitized by the ichneumonid hyperparasitoid Mesochorus lilioceriphilus Schwenke. Further details of the biology of the parasitoids are described, and their potential as biological control agents is discussed.     

Fluctuating asymmetry and developmental instability in evolutionary biology: past, present and future

The role of developmental instability (DI), as measured by fluctuating asymmetry (FA), in evolutionary biology has been the focus of a wealth of research for more than half a century. In spite of this long period and many published papers, our current state of knowledge reviewed here only allows us to conclude that patterns are heterogeneous and that very little is known about the underlying causes of this heterogeneity. In addition, the statistical properties of FA as a measure of DI are only poorly grasped because of a general lack of understanding of the underlying mechanisms that drive DI. If we want to avoid that this area of research becomes abandoned, more efforts should be made to understand the observed heterogeneity, and attempts should be made to develop a unifying statistical protocol. More specifically, and perhaps most importantly, it is argued here that more attention should be paid to the usefulness of FA as a measure of DI since many factors might blur this relationship. Furthermore, the genetic architecture, associations with fitness and the importance of compensatory growth should be investigated under a variety of stress situations. In addition, more focus should be directed to the underlying mechanisms of DI as well as how these processes map to the observable phenotype. These insights could yield more efficient statistical models and a unified approach to the analysis of patterns in FA and DI. The study of both DI and canalization is indispensable to obtain better insights in their possible common origin, especially because both have been suggested to play a role in both micro- and macro-evolutionary processes.     

Biology of the stem sawflies Trachelus tabidus and Cephus pygmaeus in the Negev of southern Israel

Wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.) stems in the Negev Desert of southern Israel are subject to mass infestations by stem sawflies larvae, primarily Trachelus tabidus F. (Cephidae, Hymenoptera). Infestations up to 55% and 50% in wheat and barley, respectively, were found in 1982/83. In addition, up to 8.6% of wild oat (Avena sp.) stems were cut by sawfly larvae in June 1983.The phenology of the insects was studied in 1981 to 1983 using yellow watertraps to catch adults and by dissecting infested stems to establish the pre-imaginal stages. Adults emerged at the end of February at magen and at the beginning of March at Gilat. Eggs and newly emerged larvae appeared in April. Larvae in diapause were found in cut stubs at the end of April and beginning of May. Pupae were found at the end of January and February.Of the sawfly specimens trapped in 1982 and 1983, the great majority was identified as T. tabidus and less than 5% as Cephus pygmaeus (Cephidae, Hymenoptera). The males of the sawflies appeared 4–6 days before the females, and were less numerous than them.About 12% of the larvae were parasitised by Tetrastichus sp. (Chalcidoidea, Eulophidae) in wheat, barley and wild oat, and about 4% by Collyria sp. (Ichneumoidea, Ichneumonidae) in wheat and barley stubs.

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Résumé Les tiges de blé et d'orge subissent des attaques massives de tenthrèdes dans le désert du Négev au Sud d'Israël. Des attaques atteignant 55 et 50,4% ont été observée respectivement sur blé et orge en 1982/83.De plus, jusqu'à 8,6% de tiges d'avoine sauvage ont été coupées par les larves de tenthrèdes en juin 1983.La phénologie des insectes a été étudiée de 1981 à 83 en utilisant des pièges jaunes pleins d'eau pour capturer les adultes et en disséquant les tiges attaquées pour déterminer les stades préimaginaux.Les adultes ont émergé à fin de février et au début mars. Les oeufs et les larves sont apparus en avril. Les larves en diapause ont été trouvées dans les souches coupées à la fin de janvier et en février.La grande majorité de Cephidae adultes piégés en 1982 et 1983 a été identifiée Trachelus tabidus F., Cephus pygmaeus L. correspondait à moins de 5%. Les mâles de tenthrèdes sont apparus 4 à 6 jours avant les femelles qui étaient plus nombreuses.Environ 12% des larves étaient parasitées par un eulophide (Tetrastichus sp.) sur blé, orge et avoine sauvage, tandis que 3,6% l'étainent par un ichneumonide (Collyria sp.) dans les souches de blé et d'orge.

     

Fluctuating asymmetry in a secondary sexual trait: no associations with individual fitness, environmental stress or inbreeding, and no heritability

It has been suggested that fluctuating asymmetry (FA) in secondary sexual traits may be a useful indicator of either individual quality or environmental stress. We tested this concept using a series of analyses of FA in male antler size in a wild red deer (Cervus elaphus) population, using four measures of size repeated across successive years on the same individuals. We found no consistent evidence of correlations between traits in levels of FA, nor of any associations between known environmental or developmental conditions. None of the four measures of FA showed a significant heritability (average h2 = 0.041), nor was there any evidence of inbreeding depression. For three of the four traits, fluctuating asymmetry did not predict either annual or lifetime breeding success. However there were significant associations between breeding success and FA in antler length. Given the series of null results in our other tests, it seems likely that this was a direct mechanistic effect rather than because measures of FA were indicative of individual quality or condition.     

Fluctuating Selection,Egg Banks and Population Genetic Structure in Cyclically Parthenogenetic Species

Associations between neutral genetic markers and genes under selection have been suggested to explain the population genetic structure of neutral genes in cyclically parthenogenetic freshwater invertebrates. A simulation model was constructed in order to analyse the extrapolated consequences of observed fluctuations in genotype frequencies in Daphnia, in the presence of egg banks. When of sufficient depth and magnitude, egg banks in combination with fluctuating selection were shown to maintain genetic variation within populations indefinitely. The level of equilibrium diversity increased with the depth and magnitude of the banks, and with intensity of selection. The same threshold was responsible for genetic differentiation between populations, which was independent of migration rate, and which was attained very rapidly following initial Hardy–Weinberg equilibrium. In the absence of selection, egg banks increased the effective size of local populations, thereby decreasing genetic differentiation at migration-drift equilibrium. These results suggest that egg banks are crucial to the genetic structure in the presence of fluctuating selective pressures, but more data are needed if this knowledge is to be used in an improved general understanding of the genetic structure of cyclically parthenogenetic species.