An experimental examination of the effects of habitat quality on the dispersal and local abundance of the butterfly Parnassius smintheus

Abstract 1. Nectar flower abundance was manipulated through flower removal, and sex ratio was manipulated by moving individual butterflies within a series of nine alpine meadows. The movement and abundance of the butterfly Parnassius smintheus in the meadows were monitored using mark–release–recapture methods.
2. A total of 937 butterflies, 698 males and 239 females, was captured. There were 223 observed between-meadow movements. Fifty-two per cent of males and 35% of females moved among meadows.
3. The immigration of male butterflies was related positively to nectar flowers, host plant abundance, and female butterflies. Male emigration was not affected by any of the treatments. The number of males captured was related positively to nectar flowers and host plants but not affected by sex ratio. The number of resident male butterflies was greater in meadows containing flowers and was related positively to host plant abundance, but unaffected by sex ratio.
4. Flower removal, sex ratio, and abundance of Sedum had no significant effect on the abundance, movement, or residence time for female butterflies, in part due to small sample size.
5. The fact that males immigrate to higher quality meadows suggests that male butterflies are assessing meadow quality, either by sampling meadows or potentially from a distance using olfactory cues.     

Effects of experimental population extinction for the spatial population dynamics of the butterfly Parnassius smintheus

While many studies have examined factors potentially impacting the rate of local population extinction, few experimental studies have examined the consequences of extinction for spatial population dynamics. Here we report results from a large‐scale, long‐term experiment examining the effects of local population extinction for the dynamics of surrounding populations. From 2001–2008 we removed all adult butterflies from two large, neighboring populations within a system of 17 subpopulations of the Rocky Mountain Apollo butterfly, Parnassius smintheus. Surrounding populations were monitored using individual, mark–recapture methods. We found that population removal decreased immigration to surrounding populations in proportion to their connectivity to the removed populations. Correspondingly, within‐generation population abundance declined. Despite these effects, we saw little consistent impact between generations. The extinction rates of surrounding populations were unaffected and local population growth was not consistently reduced by the lack of immigration. The broader results show that immigration affects local abundance within generations, but dynamics are mediated by density‐dependence within populations and by broader density‐independent factors acting between generations. The loss of immigrants resulting from extinction has little impact on the persistence of local populations in this system.     

Both spatiotemporal connectivity and habitat quality limit the immigration of forest plants into wooded corridors

Extensive afforestation of agricultural areas has increased the importance of green corridors as a dispersal network. We tested the effect of spatiotemporal connectivity, edge effect and habitat structural quality of wooded corridors on the long-term immigration success of forest specialist plants relative to the performance of forest generalists. In agricultural landscapes of central and southern Estonia, we sampled 28 historically connected and 52 isolated tree lines and alleys with a minimum age of 50 years, and 93 edges of ancient forests. The regional pool of common forest plants was compiled using species’ frequency data in 91 ancient forests. Both landscape connectivity and habitat quality affected the richness of response groups, but specialists and generalists responded to different drivers. Forest specialists required long-term neighbourhoods of ancient forest and benefited from a direct connection between forest and corridor. Habitat generalists reacted positively to the recently modified structure of the landscape. When a corridor was connected to forest, the dual edge in the corridor did not result in an increased negative edge effect on forest specialist arrival. Even if both specialists and generalists required wide corridors with optimum shade, forest specialists also benefited from mature overstorey and outward overhanging branches, whereas forest generalists used disturbance-created microhabitats. We conclude that only wooded corridors with long-term connectivity to seed source forests and widely branched tree canopies will function as a green infrastructure supporting forest-specific biodiversity.     

Management affects the pollinator abundance but not the reproductive success of butterfly orchids

Management of semi-natural grasslands should be based on the requirements of plants as well as their pollinators since conditions beneficial for plants are not necessarily beneficial for their pollinators and vice versa. The factors affecting the reproductive success (fruit set) of Platanthera bifolia and Platanthera chlorantha and their pollinators in agricultural landscape and woodlands were studied. In the years 2014–2016, we observed and caught moths during the flowering period of Platanthera species (late June–mid July) in four pure P. bifolia, five pure P. chlorantha and nine mixed populations under management or no management in Estonia. We determined pollinator species richness, pollinator abundance, fruit set and visibility of Platanthera plants in each population. We found that pollinator assemblages of P. bifolia and P. chlorantha did not differ between managed and unmanaged sites. Pollinator abundance had an effect on the fruit set of P. bifolia but not on that of P. chlorantha. Presence or absence of management, visibility of plants and rarefied pollinator species richness did not affect the fruit set of either plant species. Pollinator abundance was significantly higher in unmanaged populations of both plant species but rarefied pollinator species richness was higher only in unmanaged populations of P. chlorantha. Based on our findings the recommendations for management of semi-natural grasslands are to promote larger landscape diversity for the benefit of moth abundance by leaving unmanaged patches in different parts of a grassland in different years and rotational and post-fruiting management for higher reproductive success of orchids.     

Experimental habitat fragmentation increases linkage disequilibrium but does not affect genetic diversity or population structure in the Amazonian liverwort Radula flaccida

Habitat fragmentation increases the migration distances among remnant populations, and is predicted to play a significant role in altering both demographic and genetic processes. Nevertheless, few studies have evaluated the genetic consequences of habitat fragmentation in light of information about population dynamics in the same set of organisms. In a 10,000-km(2) experimentally fragmented landscape of rainforest reserves in central Amazonia, we examine patterns of genetic variation (amplified fragment length polymorphisms, AFLPs) in the epiphyllous (e.g. leaf-inhabiting) liverwort Radula flaccida Gott. Previous demographic work indicates that colonization rates in this species are significantly reduced in small forest reserves. We scored 113 polymorphic loci in 86 individuals representing five fragmented and five experimentally unmanipulated populations. Most of the variation (82%) in all populations was harboured at the smallest (400 m(2)) sampling unit. The mean ((+/-) SD) within-population genetic diversity (Nei's), of forest remnants (0.412 +/- 0.2) was indistinguishable from continuous (0.413 +/- 0.2) forests. Similarly, F(ST) was identical among small (1- and 10-ha) and large (> or = 100-ha) reserves (0.19 and 0.18, respectively), but linkage disequilibrium between pairs of loci was significantly elevated in fragmented populations relative to those in continuous forests. These results illustrate that inferences regarding the long-term viability of fragmented populations based on neutral marker data alone must be viewed with caution, and underscore the importance of jointly evaluating information on both genetic structure and demography. Second, multilocus analyses may be more sensitive to the effects of fragmentation in the short term, although the effects of increasing linkage disequilibrium on population viability remain uncertain.     

An assessment of direct and indirect effects of climate change for populations of the Rocky Mountain Apollo butterfly (Parnassius smintheus Doubleday)

Abstract Climate change is occurring and insects are responding. Current challenges for ecologists and managers are predicting how organisms will respond to continuing climate change and determining how to mitigate potential negative effects. In contrast to broad scale predictions for climate change involving the distribution of species, in this article we highlight the many ways in which local populations of the Rocky Mountain Apollo butterfly (Parnassius smintheus Doubleday) are predicted to respond to climate change. Using experimental and observational data collected over the past 15 years, we detail both direct and indirect effects. In addition, we identify limitations in our knowledge restricting the ability to predict how populations will respond to climate change. Some changes, such as warmer winter temperatures, may have beneficial effects; however, most of the effects of climate change will be detrimental. Variability in snow cover during the overwintering period and habitat loss due to forest encroachment have the largest potential negative effects.     

Potential metapopulation structure and the effects of habitat quality on population size of the endangered False Ringlet butterfly

The False Ringlet (Coenonympha oedippus) is a European butterfly species, endangered due to the severe loss and fragmentation of its habitat. In Hungary, two remaining populations of the butterfly occur in lowland Purple Moorgrass meadows. We studied a metapopulation occupying twelve habitat patches in Central Hungary. Our aim was to reveal what measures of habitat quality affect population size and density of this metapopulation, estimate dispersal parameters and describe phenology of subpopulations. Local population sizes and dispersal parameters were estimated from an extensive mark–release–recapture dataset, while habitat quality was characterized by groundwater level, cover of grass tussocks, bush cover, height of vegetation and grass litter at each habitat patch. The estimated size of the metapopulation was more than 3,000 individuals. We estimated a low dispersal capacity, especially for females, indicating a very low probability of (re)colonization. Butterfly abundance and density in local populations increased with higher grass litter, lower groundwater level and larger area covered by tussocks. We suppose that these environmental factors affect butterfly abundance by determining the microclimatic conditions for both larvae and adult butterflies. Our results suggest that the long-term preservation of the studied metapopulation needs the maintenance of high quality habitat patches by appropriate mowing regime and water regulation. Management also should facilitate dispersal to strengthen metapopulation structure with creating stepping-stones or gradually increase habitat quality in present matrix.     

Nesting season,nest age,and disturbance,but not habitat characteristics,affect nest survival of Chinese grouse

Nest survival is a vital component of breeding success,and affects population dynamics,as the loss of nests is the main cause of reproductive failure in birds.To identify key factors for the conservation of Chinese grouse Tetrastes sewerzowi,we tested the effects of nest concealment,nest age,nesting season,and habitat edge on nest daily survival rate(DSR)of Chinese grouse using 54 nests found at Lianhuashan Nature Reserve,Gansu,China,2009-2012.Moreover,we controlled for the effect of research activity by testing the effect of nest checks on DSR.Overall,mammal predation caused 93%of nest failures.DSR was 0.986±0.0038 in the constant model and the probability of a nest with a full clutch of 6 eggs surviving the entire 40-day nesting period was 0.526±0.090.DSR decreased with nest age and nesting season(from 19 May to 3 July).Mammals instead of avian predators being responsible for most nest failures suggest that nest sites might be selected to avoid visual avian predators,but not olfactory mammalian predators,and the decreasing trend of DSR with nest age and nesting season could attribute to an additive exposure effect.Moreover,nest checks conducted by investigators significantly lowered nest DSR,especially during the late period of nesting season and for older nests.Mammalian predators might locate the nest site by following the investigator's odor.Based on our results,we suggest that the late incubation stage is a particularly vulnerable period for nest survival of Chinese grouse and those researchers should adjust their activities around nests to balance the need of acquiring accurate data and decreasing nest predation risk.     

The bionomics,habitat requirements and population threats of the butterfly Bhutanitis thaidina in Taibai Mountain

Bhutanitis thaidina Blanchard is a rare and endangered species in China. Based on 2 years of field surveys in the town of Houzhenzi on the south slope of Taibai Mountain, this paper presents documented knowledge of this butterfly’s life history and behaviors, and indicates for the first time the reasons this butterfly is endangered in the field. Life tables show that the L1–L3 larval periods experience a high mortality. Natural enemies (mainly spiders and ants) and unfavorable climatic conditions are mainly important threats for the larval stages. Interspecific competition is also a potential threat to B. thaidina. The host plant Aristolochia manshuriensis occurs from 1,755 to 2,280 m. The suitable habitats for B. thaidina are located at open glades with a canopy cover of 0.4–0.6 in the forests. On the basis of the low population density and scarce suitable habitats, we hypothesize that B. thaidina lives in small extinction-prone populations and survives in the isolated patches in the forests on the south slope of Taibai Mountain. The suggested conservation for long-term survival of B. thaidina is the specific habitat management such as cutting shrubs and preserving open glades in dense forest.     

Increased abundance of the red admiral butterfly Vanessa atalanta in Britain: the roles of immigration, overwintering and breeding within the country

The migratory butterfly Vanessa atalanta increased in abundance at monitored sites in Britain from 1976 to 1996. Three possible causes of the increase are improved winter survival within Britain, greater breeding success within Britain, and increased immigration.
   Trends during most of the season were similar to those of immigrant or overwintered individuals in spring; thus the evidence does not support greater breeding success in Britain. As abundance in spring was not correlated with abundance in the previous autumn, when trend was taken into account, it seemed unlikely that overwintering in Britain was important. Thus the increase in abundance was probably due to increased immigration. Incidental to the main study, the mean index per site per year was closely correlated with the collated index, the usual measure of annual fluctuations. This agreement suggests that the mean index may be a useful check for trends in monitoring data for other wide-ranging organisms.     

The density of empty shells,not habitat cover or quality,predicts the density of live Rhagada snails

Camaenid land snails are often the focus of targeted biodiversity surveys in Australia, but can be difficult to locate because they hibernate for most of the year. Searches would be more effective if the density of live snails could be predicted from features of the habitat, or from the abundance of empty shells from long dead individuals. In 53 populations of Rhagada, the dominant genus of land snail in the Pilbara bioregion of Western Australia, the density of live snails is not correlated with the cover or height of spinifex (Triodia), the main habitat plant occupied by Rhagada. On the other hand, the density of empty shells on the soil surface does explain snail population density, but the relationship is weak (r²=0.209). Nevertheless, the data suggest that searches for live Rhagada are more likely to succeed in areas where empty shells are most abundant.     

Diversity and abundance of birds in relation to forest fragmentation,habitat quality and heterogeneity

This study investigated the importance of habitat quality and habitat heterogeneity for the abundance and diversity of breeding birds in continuous forest and in forest fragments surrounded by farmland in central Sweden. Positive correlations were found between species number and area, volume of Aspen Populus tremula and habitat heterogeneity. Spatial segregation of habitats at a relatively fine-grained scale is suggested to allow for the co-occurrence of more species. The abundance of at least 18 of the species in this study was influenced by fragmentation, and nine of these species preferred fragments to forest sites. The total density of birds was higher in fragments than in forest sites, probably because several fragment species forage in farmland surrounding the sites and a few also forage at edges. Nine species were more common in forest sites than in fragments, but only one species was restricted to continuous forest. However, several fragments were relatively close to forests (150 m) and forest was common in larger scale contexts. The abundance of most species (25 of 33 species) in this study was correlated with habitat quality variables (i.e. variables measuring the size, volume and diversity of ‘tree species’). Among these habitat variables the most important was the occurrence of deciduous trees which seemed to be important for 14 species. The second most important habitat factor seemed to be the diameter of trees, which was positively correlated with the abundance of eight species of which five are hole-nesters. Among coniferous trees, six species were positively correlated with the volume of Norway Spruce Picea abies, whereas no species seemed to be correlated with the volume of Pine Pinus sylvestris.     

Different photoperiods affect proliferation of lymphocytes but not expression of cellular,humoral, or innate immunity in hamsters

In seasonal mammals, photoperiod change is associated with a suite of alterations in physiology. It has recently been proposed that the immune response is one of the systems regulated by changes in photoperiod, although this hypothesis has not been rigorously challenged by assays of functional immune responses. The aim of this study was to test the hypothesis that photoperiod modulates immune responsiveness in Syrian (Mesocricetus auratus) and Siberian (Phodopus sungorus) hamsters. Consistent with previously reported data, short-day-housed (SD) animals exhibited a significant increase in lymph node cell (LNC) numbers and increased cellular proliferation in response to the polyclonal mitogen concanavalin A compared to long-day-housed (LD) animals. In contrast, LNC numbers from intact or gonadectomized SD animals that had been sensitized with the antigen dinitrofluorobenzene (DNFB) exhibited a reduced ex vivo proliferative response and reduced production of interleukin-6 (IL-6) compared to LD animals. In vivo studies of the contact hypersensitivity response of animals that had previously been sensitized, and subsequently challenged, with DNFB were similar in SD and LD animals, as was the proliferative activity of LNC recovered from these animals. There were also no photoperiodic differences in the antidinitrophenyl antibody response of animals sensitized with DNFB, or the anti-sheep red blood cell (srbc) response of animals immunized with srbc. Furthermore, no differences could be detected in the activity of natural killer cells from spleens of LD and SD Siberian hamsters, or in lipopolysaccharide-induced IL-6 production by LD and SD Syrian hamsters in vivo. Thus, although photoperiod is able to influence factors regulating the gross number and non-antigen-specific proliferation of lymphocytes in seasonally breeding mammals, day length does not directly influence activation of an effective immune response. The authors conclude, therefore, that expression of the immune response is not directly modified or compromised by photoperiod in these seasonally breeding hamster species.     

Fish composition,but not richness or abundance,differ among Phragmites,Typha, and Schoenoplectus zones during a high-water year

Hydrobiologia - Phragmites australis (ssp australis) is an aggressive colony-forming species that displaces emergent vegetation in North America, including native bulrushes (Schoenoplectus sp.) and...     

Relative importance of resource quantity, isolation and habitat quality for landscape distribution of a monophagous butterfly

Fragmentation of food resources is a major cause of species extinction. We tested the effects of habitat area, isolation and quality for the occurrence and population density of the endangered butterfly Polyommatus coridon . Polyommatus coridon larvae are monophagous on the plant Hippocrepis comosa, and both species are specialised on calcareous grassland, which is an endangered and highly fragmented habitat type in Germany.
In 2001 we surveyed all known calcareous grasslands (n=298) around the city of Göttingen (Germany) to map the population size of H. comosa in these habitats. Further, habitat isolation (between-patch distance: 70–7220 m) and habitat quality (cover of flowering plants, height of herb layer, percent bare ground, cover of shrub layer, wind protection, inclination) were quantified. Hippocrepis comosa occurred on only 124 fragments, which were then surveyed by 20 min transect counts for adult P. coridon in 2001 and 2002.
Occurrence and population density of P. coridon were largely determined by the population size of its larval food plant, which was correlated with grassland area. Effects of habitat isolation and habitat quality on P. coridon populations contributed only little to the explanation.
In conclusion, this monophagous habitat specialist depended on large habitats with large food plant populations to exist in viable populations. Habitat isolation and quality appear to contribute to occurrence and density patterns only in landscapes where these factors shift towards extremes, therefore general recommendations for conservation programs are difficult as they depend on regional distinctions.     

The relationship between tooth wear, habitat quality and late-life reproduction in a wild red deer population

1. Molar tooth wear is considered an important proximate mechanism driving patterns of senescence in ungulates but few studies have investigated the causes of variation in molar wear or their consequences for reproductive success. 2. In this study, we assessed molar tooth wear at death among red deer Cervus elaphus of known age on the Isle of Rum, Scotland. 3. First molar height showed a decelerating decline with age. In females, the rates of molar wear with age varied with location of home range and individuals experiencing low resource competition showed reduced molar wear. We suggest that this spatial variation in molar wear is related to differences in the availability of high-quality grazing habitat and levels of resource competition. 4. There was no evidence that females with more heavily worn molars had reduced reproductive performance late in life or that first molar height was associated with reproductive senescence.     

When habitat management can be a bad thing: effects of habitat quality,isolation and climate on a declining grassland butterfly

The conservation of most temperate grassland habitats and their characteristic fauna and flora requires regular low-intensive forms of land-use to counteract natural succession. Although many species tolerate moderate disturbance regimes, some are known to be susceptible to grazing or mowing, thereby causing a management dilemma. One of these species is the Woodland Ringlet butterfly, Erebia medusa. In this study, we analysed which environmental factors determine the occurrence of E. medusa in the Diemel Valley (Central Germany). Furthermore, we conducted microclimatic measurements during the winter months to investigate the role of the litter layer as a microclimatic buffer. Patch occupancy in the Diemel Valley was well explained by the amount of litter present in a patch and connectivity to other inhabited patches. The role of local climatic conditions could not be clarified, due to inter-correlations with connectivity. During the winter, the air temperature inside the litter layer was significantly less variable than above it. We conclude that the current distribution of E. medusa in the Diemel Valley is caused by the combined effect of habitat quality and connectivity, and perhaps also by climatic factors. The importance of the litter layer reflects the dependence of E. medusa on low-intensive or absent land-use. In addition, the litter layer possibly constitutes an essential habitat element, as it buffers temperature fluctuations and thus probably reduces the energy consumption of overwintering larvae. Given the species’ preference for abandoned grasslands, the conservation of E. medusa requires a low-intensity habitat management, for example, by rotational grazing or mowing of small parts of the sites. On the landscape level, the preservation of well-connected habitat networks is important.     

Relationship between abundance, growth, egg size and fecundity in a landlocked population of longfin smelt, Spirinchus thaleichthys

Sixteen year-classes of longfin smelt, Spirinchus thaleichthys , show a consistent pattern of alternating year-class strength between even (strong) and odd (weak) years. A recent (1989) odd year-class grew as well as, if not better than, the late 1960s odd year-classes. However, fish in the even year-classes (1988 and 1990) were significantly smaller in size then the 1966 and 1968 cohorts. Associated with this reduction in growth of the even year-class fish is a decline in fecundity. Fecundity of the odd year-classes of smelt was also smaller than in the early 1960s. Mean egg diameter and gonadosomatic index were not statistically different between 1988 and 1989 year-classes but the relative fecundity was higher for the 1988 year-class than for 1989, suggesting a higher reproductive investment in the even year-class. Reduction in growth and associated reproductive traits is considered to result from a reduction in the abundance of mysids, a preferred prey species.