Strong dispersal in a parasitoid wasp overwhelms habitat fragmentation and host population dynamics

The population dynamics of a parasite depend on species traits, host dynamics and the environment. Those dynamics are reflected in the genetic structure of the population. Habitat fragmentation has a greater impact on parasites than on their hosts because resource distribution is increasingly fragmented for species at higher trophic levels. This could lead to either more or less genetic structure than the host, depending on the relative dispersal rates of species. We examined the spatial genetic structure of the parasitoid wasp Hyposoter horticola, and how it was influenced by dispersal, host population dynamics and habitat fragmentation. The host, the Glanville fritillary butterfly, lives as a metapopulation in a fragmented landscape in the Åland Islands, Finland. We collected wasps throughout the 50 by 70 km archipelago and determined the genetic diversity, spatial population structure and genetic differentiation using 14 neutral DNA microsatellite loci. We compared the genetic structure of the wasp with that of the host butterfly using published genetic data collected over the shared landscape. Using maternity assignment, we also identified full‐siblings among the sampled parasitoids to estimate the dispersal range of individual females. We found that because the parasitoid is dispersive, it has low genetic structure, is not very sensitive to habitat fragmentation and has less spatial genetic structure than its butterfly host. The wasp is sensitive to regional rather than local host dynamics, and there is a geographic mosaic landscape for antagonistic co‐evolution of host resistance and parasite virulence.     

Causes and consequences of small population size for a specialist parasitoid wasp

The parasitoid wasp Cotesia melitaearum lives in extremely small extinction-prone populations in the Åland islands of southwest Finland. Intensive observational data from two generations, a laboratory competition experiment, and 8 years of survey data were used to measure the causes, extent and consequences of small population size for this parasitoid. In the spring generations of 1999 and of 2000 we observed 21 out of 23 and 26 populations respectively, ranging in size from 2 to 103 parasitoid cocoons. Within these populations the fraction of individuals surviving to adulthood decreased with increasing parasitoid population size. The largest source of mortality was predation (44%) followed by parasitism (20%) and unknown causes (10%). In the field about 30% of the host butterfly larvae are parasitized by a competing parasitoid, Hyposoter horticola. A laboratory competition experiment showed that C. melitaearum eggs died when laid in post-diapause host larvae occupied by H. horticola. Consequently one-third of the progeny of the over-wintering generation of C. melitaearum from the field die as a result of larval competition. The survey of host and parasitoid population dynamics over 8 years showed that extinction of local host butterfly populations occupied by the parasitoid was not associated with current parasitoid population size. Over the same period small parasitoid populations were more likely to become extinct than large populations. However, parasitoid population size was not related to parasitoid extinction when the host also became extinct. These data suggest that the parasitoid populations are kept small through the action of natural enemies and competitors, some of which are density dependent. Local populations are so small that they become extinct frequently and rarely measurably affect the population dynamics of their host. It is likely that this parasitoid persists in Åland because of the spatial asynchrony of local population dynamics.     

Characterization of microsatellite loci in two primary parasitoids of the butterfly Melitaea cinxia,Cotesia melitaearum and Hyposoter horticola (Hymenoptera)

Five polymorphic microsatellite loci were isolated from Cotesia melitaearum (Wilkinson) and three from Hyposoter horticola (Gravenhost). Both of these parasitic Hymenoptera use the butterfly Melitaea cinxia (Linnaeus) exclusively as their host in the Åland islands in SW Finland. Allelic diversity and heterozygosity were quantified in samples from the Åland islands and other localities around Europe. In addition, microsatellite loci for C. melitaearum were tested in five other Cotesia species: C. acuminata (Reinhard), C. bignellii (Marshall), C. congregata (Say), C. cynthiae (Nixon) and C. koebelei (Riley) from several different host species in Europe and Asia.     

Oviposition and reproductive performance of a generalist parasitoid (Trichogramma pretiosum) exposed to host species that differ in their physical characteristics

The response of generalist egg parasitoids to alternative natural hosts that are present simultaneously is not well known. We investigated the behavior of Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) in relation to two field hosts Helicoverpa armigera Hübner and Spodoptera litura Fabricius, in choice and no choice tests. We quantified the effects of natal host species and post-emergence adult age on the oviposition preference of the parasitoids. H. armigera eggs were consistently preferred over S. litura eggs, regardless of the natal host and adult age. When only S. litura eggs were available as hosts, they were parasitized at statistically similar rates to H. armigera eggs (average of 17 ± 2.7 vs. 13 ± 3.0, H. armigera to S. litura). The adult lifespan and lifetime fecundity of T. pretiosum were variable but were affected by natal host species and/or host species to which they were exposed. Mean lifespan and fecundity of parasitoids that had developed in H. armigera eggs and were exposed to H. armigera eggs for oviposition were 13.9 ± 1.8 days and 98.7 ± 11.0 adult offspring. By contrast, those that developed in S. litura eggs and were exposed to S. litura eggs for oviposition lived for 7 ± 0.9 days and produced 53.8 ± 8.0 adult offspring. The ovigeny index (OI) was significantly lower in the parasitoids exposed to H. armigera eggs than in those exposed to S. litura eggs, regardless of the natal host, indicating that H. armigera eggs sustain the adult parasitoids better than S. litura eggs. These results are used to predict parasitoid behavior in the field when both hosts are available.     

Local and spatial dynamics of a host–parasitoid system in a field experiment

Local extinction and colonisation rates are key factors in host–parasitoid metapopulation theory, but experimental evidence from the field is scarce. We studied the host–parasitoid system consisting of the aphid Metopeurum fuscoviride and its specialist parasitoid Lysiphlebus hirticornis. This system is characterised by a patchy distribution of the host plants (Tanacetum vulgare) and by frequent extinctions of local aphid populations. In a first field experiment, we found that the presence of the parasitoid increases the likelihood of extinction of local host populations (=all aphids living on one plant). In a second field experiment, we manipulated the distance between local host populations. Parasitoid colonisation rate strongly decreased with increasing distance between local host populations. Thus, our results show the importance of parasitoids for local host populations extinction and of distance between local host populations for parasitoid colonisation rate, suggesting the importance of spatial processes for host–parasitoid systems in the field.     

Variation in demographic and fine-scale genetic structure with population-history stage of <Emphasis Type="Italic">Hemerocallis taeanensis</Emphasis> (Liliaceae) across the landscape

Fine-scale genetic structure (FSGS) in plant populations is expected to be influenced by variation in demographic processes across space and over time. I chose Hemerocallis taeanensis (Liliaceae), a perennial herb with a rapid population turnover, to quantify how demographic structure and FSGS change with a population’s history (i.e., density). Nonaccumulative O-ring statistic and spatial autocorrelation analysis (kinship coefficient, F _ij ) were used to quantify spatial patterns of individuals and FSGS in four populations belonging to two population stages (expansion and maturation) in west-central Korea. The O-ring function revealed that significant aggregation of individuals occurs at short spatial scales during the earlier stage of population expansion, which reflects restricted seed dispersal around maternal individuals. However, this pattern disappears as the population density increases during population maturation, probably due to a high population density. Significant evidence of FSGS was found in two populations at the stage of population expansion (Sp, a statistic which describes the rate of decrease of pairwise kinship with distance, was 0.018 and 0.029). The results show that most seeds fall around maternal plants when initially established colonists proliferate at suitable microhabitats. In contrast to this, much lower Sp values (−0.003 and 0.004) were estimated for two populations at the stage of population maturation, which may result from the overlapping of seed shadows due to high adult density. All of these results demonstrate considerable variation in within-population demographic and genetic structures of H. taeanensis with respect to population temporal stage across the landscape.     

Genetic differentiation within metapopulations of Euphydryas aurinia and Melitaea phoebe in China

We analyzed genetic differentiation within metapopulations of two species of checkerspot butterfly, Euphydryas aurinia and Melitaea phoebe, in China. To generate genetic information, we used a new molecular technique, DALP – direct amplified length polymorphism. AMOVA results showed that most of the variation occurred among individuals within local populations of both E. aurinia and M. phoebe. However, while there was differentiation among local population in E. aurinia (P < 0.001), there was no subdivision in metapopulation of M. phoebe (P = 0.210). This is consistent with the behavior of M. phoebe adults being more dispersive than E. aurinia. Within the M. phoebe metapopulation, three neighboring patches were always occupied during the observation period (1998–2000). In addition, the number of individuals in these three populations accounted for the majority of M. phoebe larvae, and hence we conclude that the M. phoebe metapopulation might exist as a source-sink metapopulation. On the other hand, the E. aurinia metapopulation is an example of a classical metapopulation. Therefore, the conservation management of these two species should reflect these differences.     

Comparison of genetic population structure of the large blue butterflies Maculinea nausithous and M. teleius

We investigated the genetic population structure of two rare myrmecophilous lycaenid butterflies, Maculinea nausithous and M. teleius, which often live sympatrically and have similar biology. In Europe, both species occur in highly fragmented populations and are vulnerable to local extinction. The proportion of variable allozyme loci, average heterozygosity and genetic differentiation among populations was higher in M. nausithous than in sympatrically living M. teleius populations. We hypothesise that the differences in heterozygosity are mainly due to the known higher efficiency of typical host ant nests in rearing M. nausithous pupae compared to M. teleius pupae. This implies a larger probability of larval survival in M. nausithous, which buffers populations against environmental and demographic stochasticity. In contrast, the lower carrying capacity of ant nests in rearing M. teleius pupae requires higher nest-densities and makes M. teleius populations more prone to losing genetic variation through drift if this condition is not fulfilled. The single investigated Russian population of M. teleius showed much higher levels of heterozygosity than any of the Polish populations, suggesting a more viable and still intact metapopulation structure.     

The co-occurrence of an introduced biological control agent (Coleoptera: <Emphasis Type="Italic">Coccinella septempunctata</Emphasis>) and an endangered butterfly (Lepidoptera: <Emphasis Type="Italic">Lycaeides melissa samuelis</Emphasis>)

Whether a biological control agent presents a non-target risk to a native species depends if they co-occur spatially and temporally, and if the agent will harm the native species. We sampled two study sites during 1993 in Minnesota and Wisconsin to survey predators and parasitoids of the extant populations of the United States federally endangered Karner blue butterfly, Lycaeides melissa samuelis. We found the introduced coccinellid Coccinella septempuntata co-occurring spatially and temporally with eggs, larvae and adults of L. m. samuelis. The two species were also observed together on the latters sole host plant, Lupinus perennis, and in Wisconsin, an adult C. septempunctata was observed consuming second instar larvae of L. m. samuelis. Using a simple model to hypothesize the risk that C. septempunctata presents to L. m. samuelis, we showed that increases in predator density could greatly increase mortality to L. m. samuelis. At these sites, C. septempunctata were reproducing and had access to summer aphids and suitable overwintering habitat. Nearby agricultural crops could provide spring aphids for oogenesis, and assist with C. septempunctata population build-up. Maintaining a minimum isolation distance between agricultural crops known to harbor aphids and extant L. m. samuelis populations may need to be considered as part of the butterfly management program.     

Combining larval habitat quality and metapopulation structure – the key for successful management of pre-alpine Euphydryas aurinia colonies

This study aims to analyse larval habitat preferences and landscape level population structure of the threatened Marsh Fritillary butterfly, Euphydryas aurinia, and discusses implications for the conservation and management of this strongly declining species in central Europe. Whereas current management strategies are mainly based on studies of habitat requirements of adult individuals, we intend to emphasise larval habitat quality and population processes at the landscape level as additional key factors. Microhabitat preference analysis of egg-laying females showed that eggs were predominantly laid on prominent large-sized host plant individuals. Additionally, when Succisa pratensis was used as a host plant (as opposed to Gentiana asclepiadea), host individuals in open vegetation structure were preferred. Optimal oviposition conditions were present in recently abandoned calcareous fen meadows and at the edges of such meadows currently in use. A two-year patch-occupancy study in the northern pre-alpine region of south-west Germany indicated that E. aurinia lives in a metapopulation. In a logistic-regression model, patch size, isolation, and habitat quality explained 82% of the observed patch-occupancy pattern in 2001. Our data suggest that a suitable conservation strategy must incorporate both the conservation of a network of suitable habitat patches, and efforts to maximise local habitat quality by ensuring that host plants can grow to a large size and are surrounded by sparse and low vegetation cover.     

The fragmented population structure of the Sardinian chalk hill blue butterfly (Lepidoptera, Lycaenidae)

The endemic Sardinian chalk hill blue butterfly, Polyommatus coridon gennargenti, is considered vulnerable to extinction because of its low genetic variation and restricted distribution. The species also has a fragmented distribution, which follows the patchy distribution pattern of its larval host-plant. A preliminary investigation of the population structure of P. coridon gennargenti was carried out on a small network of four local populations by means of capture–recapture methods. Estimated population sizes and movement rates among the four adjacent local populations suggest that this taxon has a metapopulation structure composed of loosely connected small local populations. Natural fragmentation, isolation, and traditional land use contribute to the vulnerability of P. coridon gennargenti to extinction. Low effective population sizes and restricted movement between habitat patches lead to inbreeding and an increased vulnerability to extinction of this island population.     

Comparative population structure of Chinese sumac aphid <Emphasis Type="Italic">Schlechtendalia chinensis</Emphasis> and its primary host-plant <Emphasis Type="Italic">Rhus chinensis</Emphasis>

Most of our current understanding of comparative population structure has been come from studies of parasite–host systems, whereas the genetic comparison of gallnut-aphids and their host-plants remain poorly documented. Here, we examined the population genetic structure of the Chinese sumac aphid Schlechtendalia chinensis and its unique primary host-plant Rhus chinensis in a mountainous province in western China using inter-simple sequence repeat (ISSR) markers. Despite being sampled from a mountainous geographic range, analysis of molecular variance (AMOVA) showed that the majority of genetic variation occurred among individuals within populations of both the aphid and its host. The aphid populations were found to be structured similarly to their primary host populations (F _ST values were 0.239 for the aphid and 0.209 for its host), suggesting that there are similar patterns of gene flow between the populations of the aphid and between populations of its host-plant. The genetic distances (F _ST/1 − F _ST) between the aphid populations and between its host-plant populations were uncorrelated, indicating that sites with genetically similar host-plant populations may not always have genetically similar aphid populations. The lack of relationships between genetic and geographical distance matrices suggested that isolation by distance (IBD) played a negligible role at this level. This may be mainly attributed to the founder effect, genetic drift and the relative small spatial scale between populations. Zhumei Ren and Bin Zhu contributed equally to this work.     

Response of the wasp (<Emphasis Type="Italic">Cotesia glomerata</Emphasis>) to larvae of the large white butterfly (<Emphasis Type="Italic">Pieris brassicae</Emphasis>)

The large white butterfly (Pieris brassicae L) first invaded northernmost Japan from Siberia around 1994, and after a few years, began to expand its range. The wasp, Cotesia glomerata (L) parasitizes larvae of the small white butterfly (Pieris rapae crucivora Boisduval), a usual host in the same geographic area. Some Pieris brassicae larvae in Hokkaido have been parasitized by Cotesia glomerata, but the parasitism rate of Pieris brassicae larvae tends to be lower than that of Pieris rapae. To examine the process of parasitizing Pieris brassicae larvae, we observed how the parasitoid wasp responded to the host larvae on damaged leaves. Cotesia glomerata females tended to avoid Pieris brassicae larvae, and even when female wasps inserted their ovipositors into Pieris brassicae larvae, none laid eggs. The parasitoids obtained from Pieris rapae larvae failed to parasitize Pieris brassicae during the host-acceptance step.     

Relative importance of host plant patch geometry and habitat quality on the patterns of occupancy,extinction and density of the monophagous butterfly <Emphasis Type="Italic">Iolana iolas</Emphasis>

Habitat fragmentation is a major cause of species rarity and decline because it increases local population extinctions and reduces recolonisation rates of remnant patches. Although two major patch characteristics (area and connectivity) have been used to predict distribution patterns in fragmented landscapes, other factors can affect the occurrence of a species as well as the probability of it becoming extinct. In this paper, we study the spatial structure and dynamics of the butterfly Iolana iolas in a 75-patch network of its host plant (Colutea hispanica) to determine the relative importance of patch area, connectivity and habitat quality characteristics on occupancy, extinction and density over the period 2003–2006. Occupancy in 2003, incidence (proportion of years occupied) and probability of extinction were mostly affected by patch area. Smaller patches were less likely to be occupied because they had a higher probability of extinction, partly due to environmental stochasticity. The density of I. iolas was negatively related to patch area in all study years. Only in 2004 was the density of I. iolas positively influenced by fruit production per plant. Our results suggest that for I. iolas, and probably for other specialist butterflies with clearly delimited resource requirements, metapopulation dynamics can be satisfactorily predicted using only geometric variables because most habitat characteristics are subsumed in patch area. However, this hypothesis should be subject to further testing under diverse environmental conditions to evaluate the extent of its generalisation.     

Population Genetic Structure of Two Columnar Cacti with a Patchy Distribution in Eastern Brazil

The genetic variability and population genetic structure of six populations of Praecereus euchlorus and Pilosocereus machrisii were investigated. The genetic variability in single populations of Pilosocereus vilaboensis, Pilosocereus aureispinus, and Facheiroa squamosa was also examined. All of these cacti species have a patchy geographic distribution in which they are restricted to small areas of xeric habitats in eastern Brazil. An analysis of genetic structure was used to gain insights into the historical mechanisms responsible for the patchy distribution of P. euchlorus and P. machrisii. High genetic variability was found at the populational level in all species (P=58.9–92.8%, A_p=2.34–3.33, H_e=0.266–0.401), and did not support our expectations of low variability based on the small population size. Substantial inbreeding was detected within populations (F_IS=0.370–0.623). In agreement with their insular distribution patterns, P. euchlorus and P. machrisii had a high genetic differentiation (F_ST=0.484 and F_ST=0.281, respectively), with no evidence of isolation by distance. Accordingly, estimates of gene flow (N_m) calculated from F_ST and private alleles were below the level of N_m=1 in P. machrisii and P. euchlorus. These results favored historical fragmentation as the mechanism responsible for the patchy distribution of these two species. The genetic distance between P. machrisii and P. vilaboensis was not compatible with their taxonomic distinction, indicating a possible local speciation event in this genus, or the occurrence of introgression events.     

珍稀濒危植物黄杉种群结构与空间分布格局北大核心CSCD

为了解珍稀濒危植物黄杉种群的生存现状和更新机制,对贵州省威宁县黄杉自然保护区内天然分布的黄杉(Pseudotsuga sinensis)群落的结构特征和空间分布格局进行了研究。结果表明,黄杉种群的中龄级个体数较多,幼龄级和老龄级个体相对较少,呈典型的“∩”型结构。种群结构动态指数V_(pi)(13.49%)和V′_(pi)(0.05%)均大于0,种群属于增长型,但增长幅度逐渐降低,对外界干扰敏感度较高。种群存活曲线趋于Deevey-II型,生命期望值单调下降,死亡率呈“V”型,有2个死亡高峰,第1个出现在I龄级,第2个出现在IX龄级。黄杉种群除部分样地(Q2和Q7)呈均匀分布外,大部分样地呈集群分布;随龄级的增加,空间分布格局由集群分布逐渐趋向于均匀分布。因此,黄杉种群濒危的主要因素是幼树个体无法及时补充种群,建议加强对幼苗的抚育和通过适度干扰,促进黄杉种群的更新。     

Spatial genetic structure within two populations of a self-incompatible perennial,Chionographis japonica var.japonica (Liliaceae)

Intrapopulational spatial genetic structure was examined in two populations ofChionographis japonica var.japonica, a self-incompatible perennial, by spatial autocorrelation analysis of enzyme polymorphism. Although most spatial autocorrelation indices (Moran'sI) in the shortes distance class were significantly positive, most in the other distance classes did not significantly deviate from the values expected from random distributions of genotypes in both populations. This contrasts with a spatial genetic pattern previously reported for a population of the predominantly selfing congener,C. japonica var.kurohimensis, indicating that pollen-mediated gene flow highly impedes genetic substructuring within populations of outcrossingC. japonica var.japonica. Genetic similarity in very proximate distance found in outcrossingC. japonica var.japonica is probably due to restricted dispersal of seeds.     

Nectar exploitation by herbivores and their parasitoids is a function of flower species and relative humidity

In conservation biological control, diversification of the agro ecosystem with flowering vegetation is seen as an important tool to support the broad range of predators and parasitoids that require nectar and pollen sources to survive and reproduce. In order to identify flowering plants that provide suitable food sources for natural enemies without supporting the pest species, we analyzed the exploitation of 19 flowering plants by two important lepidopteran cabbage pests, Pieris rapae and Plutella xylostella, and their hymenopteran parasitoids, Cotesia glomerata and Diadegma semiclausum. The experiments were conducted at 90% r.h., while Pieris rapae was tested both at 45% r.h. and at 90% r.h. At 45 ± 5% r.h., corresponding with field conditions at which P. rapae is predominantly active, the butterfly was unable to feed on a number of exposed floral nectar sources whose nectar was successfully exploited at 90% r.h. The broader nectar exploitation by P. rapae at the high humidity is presumably explained by the resulting decrease in nectar viscosity. When comparing D. semiclausum and its herbivorous host P. xylostella, the herbivore exploited a broader range of plants. However, those plants that benefited both the parasitoid and the herbivore had a much stronger effect on the longevity of the parasitoid. The results from the accessibility bioassay suggest that flowers where nectar is not accessible can have a negative impact on insect survival presumably by stimulating foraging without providing accessible nectar. Our results underline the importance of considering species-specific environmental conditions when fine-tuning the choice of nectar sources to be used in conservation biological control programs.     

The effect of host size on the sex ratio of Syngaster lepidus, a parasitoid of Eucalyptus longhorned borers (Phoracantha spp.)

The solitary larval ectoparasitoid, Syngaster lepidus Brullé, parasitizes the cryptic larvae of two wood-boring beetles, Phoracantha recurva Newman and Phoracantha semipunctata F. The objective of this study was to determine how the female parasitoids allocated the sex of progeny when presented with larval hosts of uniform size classes. Host size was directly correlated with age of the Phoracantha larval hosts. Groups of Phoracantha larvae of a single age class (2-, 3-, 4-, or 5-week-old) were exposed to parasitoids, and sex ratios of the resulting parasitoid progeny from each host age class were determined. A significant relationship was observed among the sizes of P. recurva and P. semipunctata hosts and the sex ratio of emerging parasitoids. Parasitized 2-week-old beetle larvae of both Phoracantha spp. produced only male S. lepidus progeny, whereas older larval hosts produced increasing proportions of female parasitoids (up to 80% females from 5-week-old hosts). Two-week-old Phoracantha larvae of both species produced fewer parasitoids than host larvae 3–5-week-old. The size of parasitoid progeny consistently increased with host larval age (size), and female parasitoids were larger than males across all host size classes. Male S. lepidus developed in approximately 25 days from 2-week-old hosts, and 19–21 days in 3–5-week-old hosts. Female S. lepidus developed in 22–25 days, with developmental time increasing with host size.     

瓜实蝇2种蛹寄生蜂生防潜能比较

[目的] 比较研究蝇蛹俑小蜂与蝇蛹金小蜂对瓜实蝇的控制潜能。[方法] 采用非选择性实验测定蝇蛹俑小蜂与蝇蛹金小蜂对瓜实蝇蛹的寄生效能、繁殖能力,并研究覆土厚度对2种寄生蜂寄生效能的影响。[结果] 2种寄生蜂对瓜实蝇的寄生率无显著性差异,但蝇蛹金小蜂的平均单雌产后代数比蝇蛹俑小蜂的多,分别为30和23头。土壤厚度显著影响2种寄生蜂的寄生效能,随着土壤厚度的增加,2种寄生蜂对瓜实蝇蛹的寄生率均显著下降,但蝇蛹金小蜂的寄生率下降更为迅速;蝇蛹俑小蜂最深可寄生8 cm土壤下的瓜实蝇蛹,而蝇蛹金小蜂在土壤厚度达到3 cm时就不能完成寄生。[结论] 蝇蛹俑小蜂较蝇蛹金小蜂更适合应用于瓜实蝇蛹的生物防治。