Larval host plant affects fitness consequences of egg size variation in the seed beetle Stator limbatus

Egg size variation often has large effects on the fitness of progeny in insects. However, many studies have been unable to detect an advantage of developing from large eggs, suggesting that egg size variation has implications for offspring performance only under adverse conditions, such as during larval competition, periods of starvation, desiccation, or when larvae feed on low-quality resources. We test this hypothesis by examining the consequences of egg size variation for survivorship and development of a seed-feeding insect, Stator limbatus, on both a low-quality (Cercidium floridum) and a high-quality (Acacia greggii) host plant. Our results are consistent with the hypothesis. S. limbatus larval performance was affected by egg size only when developing on the poor-quality host (C. floridum); larvae from large eggs survived better on C. floridum than those from small eggs, while there was no evidence of an effect of egg size on progeny development time, body weight, or survivorship when larvae developed on A. greggii. These results indicate intense selection for large eggs within C. floridum-associated populations, but not in A. greggii-associated populations, so that egg size is predicted to vary among populations associated with different hosts. Our results also support this hypothesis; females from a C. floridum-associated population (Scottsdale) laid larger eggs than females from an A. greggii-associated population (Black Canyon City).     

Plasticity in the egg-spacing behavior of a seed beetle: Effects of host deprivation and seed patchiness (Coleoptera: Bruchidae)

Egg-laying females of the seed beetle Callosobruchus maculatusdiscriminate between egg-free and egg-laden seeds and produce a nearly uniform distribution of eggs among seeds. We examined plasticity in this trait in response to both an internal factor (level of host deprivation) and an environmental one (the spatial configuration of available seeds). Responses to each factor were measured in genetically divergent strains that show a relatively strong (S strain) or weak (B strain) tendency to spread eggs evenly among seeds. Following a modest (10-h) period of host deprivation, S-strain females distributed their eggs less uniformly among seeds; the proportion of females committing at least one oviposition mistake increased from 20 to 50%. Similarly, S-strain females distributed their eggs less uniformly if seeds were presented in multiple, discrete patches instead of in a single, large patch. The higher frequency of oviposition mistakes in the multiple-patch arena was caused in part by females maintaining a uniform distribution of eggs within patches but not among patches. In contrast, females from the sloppier B strain were unaffected by either host deprivation or resource dispersion. Responses to seed patchiness are discussed in relation to the role of learning in the egg-spacing behavior of C. maculatus.     

Population differences in host use by a seed-beetle: local adaptation, phenotypic plasticity and maternal effects

For insects that develop inside discrete hosts, both host size and host quality constrain offspring growth, influencing the evolution of body size and life history traits. Using a two-generation common garden experiment, we quantified the contribution of maternal and rearing hosts to differences in growth and life history traits between populations of the seed-feeding beetle Stator limbatus that use a large-seeded host, Acacia greggii, and a small-seeded host, Pseudosamanea guachapele. Populations differed genetically for all traits when beetles were raised in a common garden. Contrary to expectations from the local adaptation hypothesis, beetles from all populations were larger, developed faster and had higher survivorship when reared on seeds of A. greggii (the larger host), irrespective of their native host. We observed two host plant-mediated maternal effects: offspring matured sooner, regardless of their rearing host, when their mothers were reared on P. guachapele (this was not caused by an effect of rearing host on egg size), and females laid larger eggs on P. guachapele. This is the first study to document plasticity by S. limbatus in response to P. guachapele, suggesting that plasticity is an ancestral trait in S. limbatus that likely plays an important role in diet expansion. Although differences between populations in growth and life history traits are likely adaptations to their host plants, host-associated maternal effects, partly mediated by maternal egg size plasticity, influence growth and life history traits and likely play an important role in the evolution of the breadth of S. limbatus’ diet. More generally, phenotypic plasticity mediates the fitness consequences of using novel hosts, likely facilitating colonization of new hosts, but also buffering herbivores from selection post-colonization. Plasticity in response to novel versus normal hosts varied among our study populations such that disentangling the historical role of plasticity in mediating diet evolution requires the consideration of evolutionary history.     

Life-history variation in a seed beetle: adult egg-laying vs. larval competitive ability

Summary Populations of the seed beetle Callosobruchus maculatus differ genetically in several traits that mediate intraspecific competition. This study examined competitive interactions between larvae from two strains that differed in their propensity to oviposit on occupied hosts. In a strain (S) where females avoided laying >1 egg/seed, larvae were highly competitive; if two larvae entered a small host simultaneously, only one adult emerged. In a strain (I) whose females were sloppier in their egg-laying decisions, more than half of the seeds bearing two larvae yielded two adults. If seeds contained one larva from each strain, only one adult emerged per seed, and 70% of these adults belonged to the more competitive S strain. A larva's probability of emergence could be increased if it entered the seed before its competitor. A two-day headstart was needed merely for I larvae to compete equally with S larvae. Competition also affected development time and adult weight, but its effect was highly strain- and sex-specific. Adult life-history traits also differed substantially between strains. Compared to I beetles, S beetles exhibited decreased longevity, lower fecundity, a truncated period of oviposition, and larger egg and body size. Fecundity was linearly related to body size in the I strain, but was largely independent of size in the S strain. When faced with a shortage of hosts, S females (whose progeny are highly competitive) withheld eggs and died without depositing 40% of their lifetime supply. In contrast, the fecundity of I females was independent of seed availability. Comparisons with previous studies suggest that both highly uniform egg-laying and strong interference among larvae may be a consequence of chronic association with a relatively small host. Results from the larval competition experiments were used as parameter estimates for a simple, game-theoretic model that postulates interference vs. exploitative strategies. Fitness comparisons suggest that a strategy employing interference competition cannot be invaded by a more exploitative form of competition in a small host.     

Individual variation in two host plants of the ladybird beetle,Epilachna pustulosa (Coleoptera: Coccinellidae)

Individual variation in two species of host plants (thistle,Cirsium kamtschaticum, and blue cohosh,Caulophyllum robustum) of the herbivorous ladybird beetleEpilachna pustulosa was examined under laboratory conditions for their acceptability to adult beetles as a food resource, for adult preference and for larval performance. When clones of these plants were subjected to non-choice tests using posthibernating female beetles, there was found to be significant intraspecific variation among clones in terms of their acceptability, but interspecific variation was not detected. Significant intraspecific as well as interspecific variation were frequently detected in the two host plants when clones of these plants were subjected to choice tests using posthibernating female beetles; the magnitude of interspecific plant variation for beetle preference is not necessarily larger than that of intraspecific plant variation. Individual variation across plant species with respect to beetle larval performance was also significant. A positive correlation between adult preference and larval performance is suggested across the two taxonomically remote host plant species, thistle and blue cohosh, although this needs further investigation.     

Female's preference for oviposition site and larval performance in the water-lily beetle,Galerucella nymphaeae (Coleoptera: Chrysomelidae)

Water-lily beetles prefer younger rather than older water-lily leaves as oviposition sites. By the time of hatching, however, young leaves have aged consieerably. Larval performance of the water-lily beetle was measured on different types of leaves of the yellow water-lily and compared with oviposition preference of females. The leaf types used in the experiments were categorized as (i) young, (ii) natal (medium-aged) and (iii) old. The natal leaves were the ones on which larvae from a particular egg-batch had hatched. There were two sets of experiments. First, larvae were raised from eggs to pupae on young and on old leaves. Second, the growth of the 1st-instar larvae was measured on young, natal, and old leaves. The development time from egg to pupa did not differ between young and old leaves, but larvae growing on young leaves attained a higher pupal weight. In the second experiment the 1st-instar larvae grew fastest on their natal leaves, but there was also variation in the growth rate of progeny from different egg-batches. Larval growth on young and old leaves did not differ significantly. Larvae tried to emigrate much less from natal than other types of leaves. Females tended to lay eggs on leaves where larval growth was fastest. It seems that medium-aged leaves are best for larval growth, but the leaf characteristics responsible for this remain unresolved.     

Host-associated differences in fitness within and between populations of a seed beetle (Bruchidae): effects of plant variability

Summary Experiments were conducted with the sexually reproducing seed beetle Stator limbatus and its hosts in north-central Arizona to determine if it was substructured into units, each specialized for higher fitness on a specific host species. Unlike many studies, we incorporated scale, i.e., conducting experiments between and within beetle populations on seeds from within and between plant species. Of particular interest was whether intraspecific plant variability prevented beetle specialization within beetle populations. Results suggest that S. limbatus is specialized to certain hosts. On the palo verde Cercidium floridum, beetles originally reared from this host had significantly higher emergence compared to beetles transferred from other hosts. We did not test directly for a genetic basis for this. Alternative hypotheses of variation in symbiotic microorganisms in larval guts and maternal effects were assessed. Essentially no bacteria, yeast or protozoa were found, and maternal effects as expressed by varying egg weights were not detected; however, other microorganisms might have been present and maternal effects through inducible enzymes was possible. Caution, then, is needed in any genetic interpretations of our results. The differences on C. floridum were detected from tests between and within beetle populations. Evidence for specialization was not detected on the other hosts, Cercidium microphyllum and Acacia greggii. On the other hosts, beetles performed well regardless of their source. Significant differences were detected among individual plants of C. floridum as to the suitability of their seeds for deveoopment of S. limbatus. No such differences were detected among the other host plants. These patterns of conspecific plant variability are opposite of what is expected if plant variability prevents specialization of beetles to particular species of hosts. Thus, the data suggest seed variability among plants does not prevent specialization to host species in this system. We discuss how the patterns of host use in this study relate to the hypothesis of sympatric host race formation.     

Effect of experience on host preference inCallosobruchus maculatus (Coleoptera: Bruchidae): Variability among populations

Experiments using naive females established differences in host specificity among geographic strains of the seed beetleCallosobruchus maculatus; some females strongly preferred azuki bean over mung bean, and others failed to discriminate between the two hosts. We examined whether such congenital differences affect the degree to which host preference can be modified by experience. In choice tests, previous exposure to azuki bean increased the proportion of eggs laid on that host, but only in strains with a relatively low host specificity. Under more realistic, no-choice conditions, egg-laying experience affected oviposition rates mostly in strains with a high host specificity, but these experiments could not distinguish between the effects of a female's experience per se and her physiological state (i.e., egg load). Our results indicate that the likelihood of detecting an effect of experience on host choice depends on both the experimental protocol and the source of the test population. In natural populations ofC. maculatus, recent egg-laying experience probably plays little or no role in discrimination between host species but may influence discrimination between conspecific seeds that differ in quality.     

Behavioral evidence for host races in Rhagoletis pomonella flies

Summary One of the most controversial putative cases of host race formation in insects is that of the apple maggot fly, Rhagoletis pomonella (Diptera: Tephritidae). A principal cause of the controversy is lack of relevant data. In laboratory and field enclosure experiments, we compared the host acceptance behavior of sympatric populations of flies originating from naturally infested hawthorn (the native host) and apple (an introduced host) in Amherst, Massachusetts or East Lansing, Michigan. In general, hawthorn fruit were accepted for ovipositional attempts nearly equally by apple and hawthorn origin females, whereas apples were accepted much more often by apple than hawthorn origin females. Similarly, males of apple and hawthorn origin exhibited about equal duration of residence on hawthorn fruits as sites at which to acquire potential mates, while males of apple origin tended to reside substantially longer than males of hawthorn origin on apples. Irrespective of fly origin, both sexes always responded more positively to hawthorn fruit than to apples. Because all flies assayed were naive (ruling out effects of prior host experience of adults) and because tests revealed no influence of pre-imaginal fruit exposure on pattern of host fruit acceptance by females, the combined evidence suggests the phenotypic differences we observed in host response pattern between hawthorn and apple origin flies may have an underlying genetic basis. Further tests showed that while larval progeny of flies of each origin survived better in naturally growing hawthorn fruit than in naturally growing apples, there was no differential effect of fly origin on larval survival ability in either host. We discuss our findings in relation to restriction in gene flow between sympatric populations of R. pomonella and in relation to current models of host shifts in insects.     

Host plant-mediated variation in overwintering site quality: implications for the size and composition of populations of Acanthoscelides alboscutellatus (Coleoptera: Bruchidae)

This study provides an example of how variation in the quality of overwintering sites provided by the host plant of an insect seed predator can influence both the probability of overwintering survival and the size and composition of postwintering populations. Thus, the concept of host plant quality is extended to include variation in the suitability of the overwintering site of temperate region insects that overwinter within, or in habitats created by, their host plant. Adult Acanthoscelides alboscutellatus (Horn) (Coleoptera: Bruchidae) overwinter inside the fruit of Ludwigia alternifolia (L.) (Onagraceae). In early winter, however, fruits begin to dehisce, i.e., one or more of the fruit's four sides and/or top are shed. Variation in the onset and extent of dehiscence creates a range of overwintering habitats that vary in exposure to ambient conditions. In this study the frequency of possible overwintering sites in natural populations of L. alternifolia was determined by monitoring the phenology of fruit dehiscence from October through May in two populations for four years and for a third population for three years. Winter survivorship of adult A. alboscutellatus was assessed experimentally in eight environments representative of the conditions created by variation in dehiscence. These environments were produced by crossing four levels of exposure (degree of dehiscence) with two locations of the overwintering site, i.e., above or on the ground surface. The onset, phenology, and overall frequency of fruit dehiscence varied markedly among populations and years. Exposure, location, and their interaction had strong effects on survival and accounted for 80% of the observed variation in winter survival. Survivorship was higher on than above the ground, and in both locations decreased with increasing exposure. Thus, variation in fruit dehiscence among L. alternifolia populations will influence the size of postwintering A. alboscutellatus populations by dictating the quality of overwintering sites. Adult beetles that over-winter inside indehiscent fruit experience selection for small body size, associated with high mortality, when they attempt to exit the fruit at eclosion. As a consequence, the frequency of fruit dehiscence at eclosion coupled with the relative survival rates of adults within indehiscent fruit will determine the body size composition of postwintering populations and hence the response to selection for small body size in this species.     

Dietary specialization in European species groups of seed beetles (Coleoptera: Bruchidae: Bruchinae)

Because of their particular biology, seed beetles exhibit a strong relationship with their larval host plants. In Europe, however, field data have long been scarce and unreliable. The results of Legume seed collections of nearly 1,000 samples belonging to 292 species from various locations in Europe are summarized. The status of current Bruchidius species groups is amended on morphological and phylogenetic bases. Recent advances in the knowledge of phylogenetic structures of both Fabaceae and Bruchinae provide a new picture of Bruchinae–Fabaceae interactions. It reveals a certain level of host conservatism. The hypothesis of radiative adaptation seems the most compatible with observed data.According to currently accepted phylogenies, seed beetles constitute a sister group to Sagrinae within the family Chrysomelidae (see discussion in Lingafelter and Pakaluk 1997). For convenience, we have, however, retained the name Bruchidae.     

Variable correspondence of female host preference and larval performance in a phytophagous ladybird beetle Epilachna pustulosa (Coleoptera: Coccinellidae)

Genetic and environmental factors causing intraspecific variation of the thistle Cirsium kamtschaticum Ledeb. as a host plant of the phytophagous ladybird Epilachna pustulosa Kôno were investigated through simple food-choice tests and rearing of larvae. Two thistle clones (T1U₁ and T4H₂) were used, originally growing approximately 12 km apart. A previous study showed that adult female ladybirds preferred T1U₁ to T4H₂, and that larval performance was better on T1U₁, when leaves from the clones in situ were examined. The two clones retained their characteristics with respect to beetle preference after transplantation into a common garden. However, the difference between T1U₁ and T4H₂ with respect to larval performance was reduced after the transplantation. When leaves from shoots of T1U₁ exposed to different sunlight intensities were offered, adult female ladybirds did not show obvious preferences. Larval eclosion rates increased significantly with the increase in leaf sunlight intensity exposure. These results suggest strongly that both genetic and environmental factors are involved in interclonal variation of thistle quality in beetle preference and/or performance. It is suggested that the quality of thistle leaves for larval performance is largely affected by environmental factors, while leaf quality for beetle preference may be determined strictly by genetic factors. Under certain conditions, E. pustulosa females may behave maladaptively, preferring plants not appropriate for larval growth, or not choosing plants appropriate for the larval growth.     

Insights to host discrimination and host acceptance behaviour in a parasitoid (Diptera: Asilidae): Implications for fitness

The robber fly Mallophora ruficauda is one of the principal pests of apiculture in the Pampas region of Argentina. As adults they prey on honey bees and other insects, while as larvae they are solitary ectoparasitoids of third instar scarab beetle larvae. Females of M. ruficauda lay eggs away from the host in tall grasses. After being dispersed by the wind, larvae drop to the ground, where they dig in search of their hosts. It is known that second instar larvae of M. ruficauda exhibit active host searching behaviour towards its preferred host, third instar larva of Cyclocephala signaticollis. Although the means by which host location occurs has been studied and since superparasitism is a frequent scenario in the field, no information about host discrimination and host acceptance is available. We carried out studies in the field and behavioural experiments in the laboratory to determine if M. ruficauda is capable of quality host discrimination. We also studied if this parasitoid is capable of conspecific detection in order to avoid superparasitism. Finally, we analyzed the conditions under which superparasitism occurs in the field. We report here that the second instar larva of M. ruficauda is able to discriminate the parasitism status of the host by means of chemical cues, but is not capable of detecting conspecifics prior to attacking a host. We also found that the host cannot detect the presence of the parasitoid by means of chemical cues, so that no counter-defense against parasitism occurs. Furthermore, we determined that superparasitism occurs on the heavier hosts, i.e. those with more abundant resources which could harbor several parasitoid individuals. Finally, we discuss the possible implications of larval host location and host discrimination decisions on the fitness of this parasitoid.     

Intraspecific mitochondrial DNA variation in the Colorado potato beetle,Leptinotarsa decemlineata (Coleoptera: Chrysomelidae)

Restriction endonuclease analyses of mitochondrial DNA (mtDNA) were used to examine genetic variability and population structure inLeptinotarsa decemlineata (Say). A group of three enzymes, EcoRI, HpaI, and PstI, was used to reveal polymorphism both within and among some of the 10 populations tested, yielding 16 haplotypes in combination. The frequencies of these 16 haplotypes differed significantly across geographic regions, indicating some partitioning of mtDNA haplotypes. Estimates of mtDNA sequence divergence (δ) between haplotypes ranged from 0.016 to 0.135%, suggesting local differentiation of mtDNA in some populations. Analysis of these data suggests that Texas was colonized by more than one mtDNA lineage, most likely originating in Mexico. We hypothesize that a larger founder size for the initial introductions or high levels of variability in the parent population at the edge of the CPB expanding range led to the initial partitioning of haplotypes observed in samples from Texas.     

Genetic architecture underlying convergent evolution of egg-laying behavior in a seed-feeding beetle

Independent populations subjected to similar environments often exhibit convergent evolution. An unresolved question is the frequency with which such convergence reflects parallel genetic mechanisms. We examined the convergent evolution of egg-laying behavior in the seed-feeding beetle Callosobruchus maculatus. Females avoid ovipositing on seeds bearing conspecific eggs, but the degree of host discrimination varies among geographic populations. In a previous experiment, replicate lines switched from a small host to a large one evolved reduced discrimination after 40 generations. We used line crosses to determine the genetic architecture underlying this rapid response. The most parsimonious genetic models included dominance and/or epistasis for all crosses. The genetic architecture underlying reduced discrimination in two lines was not significantly different from the architecture underlying differences between geographic populations, but the architecture underlying the divergence of a third line differed from all others. We conclude that convergence of this complex trait may in some cases involve parallel genetic mechanisms.     

Consequences of inbreeding for the cowpea seed beetle, Callosobruchus maculatus (F.)(Coleoptera: Bruchidae)

Inbreeding is said to reduce vigour and fitness. It may also determine how a population responds to selection. Local populations of Callosobruchus maculatus , the cowpea seed beetle, are established annually from small numbers of founders and the species has been distributed to many parts of the world where isolated populations may have been founded by very small numbers of individuals. After more than 20 generations of inbreeding, inbred lines have been shown to diverge from a common ancestral stock in similar directions with respect of some variables such as developmental speed, but haphazardly in respect of other parameters such as male weight. The respective roles of drift and of selection as effective evolutionary forces in inbred lines are discussed in the light of these results. It is argued that some intraspecific differences in C. maculatus may be explained as a product of periodic inbreeding, but that the process does not impair the ability to adapt to local conditions so contributing to the status of the species as a pest of international importance.     

Host plant suitability of a recently naturalized thistle Cirsium vulgare (Asteraceae) for a phytophagous ladybird beetle,Epilachna pustulosa (Coleoptera: Coccinellidae)

The phytophagous ladybird beetle, Epilachna pustulosa Kôno occurs mainly on the thistle Cirsium kamtschaticum Ledeb. In recent years, we have confirmed feeding and oviposition by E.pustulosa, and some degree of larval development of this beetle, on a naturalized congeneric thistle, Cirsium vulgare (Savi) Tenore. In this study, we assessed the host suitability of C.vulgare for E.pustulosa, using larval rearing and simple food choice tests, by comparing the performance on, and preference for, their legitimate host C.kamtschaticum. All experiments were initiated in July, when the utilization of C.vulgare by E.pustulosa became obvious. For larval performance, E.pustulosa showed a lower eclosion rate, shorter developmental duration and smaller body size when reared on C.vulgare than on C.kamtschaticum. Cirsium vulgare appeared to be a less adequate larval food than C.kamtschaticum. Adult feeding preference was altered seasonally; females preferred C.kamtschaticum to C.vulgare in a food choice test conducted in early July, whereas they showed the opposite preference in tests conducted in mid and late July. The utilization of C.vulgare by E.pustulosa appeared to be induced by qualitative deterioration of C.kamtschaticum leaves in July, which resulted in increased dispersion of adult beetles.     

Post-dispersal predation of Acacia farnesiana seeds by Stator vachelliae (Bruchidae) in Central America

Summary Post-dispersal seed predation by the bruchid beetle Stator vachelliae was investigated in Santa Rosa National Park, Costa Rica. This insect finds the seeds of the leguminous Acacia farnesiana in the feces of horses, deer, and ctenosaur lizards, the current major dispersers. Patterns of oviposition and pre-adult survival of beetles in the seeds were investigated in a series of experiments using fresh horse dung. S. vachelliae never minded into the dung balls, attacking only those seeds located on the surface. Fresh horse dung did not attract insects more readily than dry dung. The proportion of seeds attacked was not related to their density in a defecation, and was similar in three areas with different densities of the host plant. In a fourth area with no fruiting A. farnesiana shrubs all seeds survived insect predation. Bruchids attacked a greater proportion of seeds at 1 m than at 5 m from the edge of the shrub's crown. Seeds were mainly removed from horse dung by rodents with similar intensity in all areas and at both distances; this seed removal interfered with bruchid oviposition and probably with bruchid survival. S. vachelliae oviposited less frequently on seeds in dung fully exposed to sun. When oviposition on a dung pile was high, the distribution of eggs on the seeds was clumped, suggesting that some seeds were preferred to others. By the end of the dry season, bruchids stopped attacking the seeds. The results show that the fate of both seeds and bruchids is greatly influenced by the location and time of defecation.     

Host adaptation in the anther smut fungus Ustilago violacea (Microbotryum violaceum): infection success,spore production and alteration of floral traits on two host species and their F1-hybrid

It is often assumed that host specialization is promoted by trade-offs in the performance of parasites on different host species, but experimental evidence for such trade-offs is scant. We studied differences in performance among strains of the anther smut fungus Ustilago violacea from two closely related host plant species, Silene alba and S. dioica, on progeny of (1) the host species from which they originated, (2) the alternative host species, and (3) inter-specific hybrids. Significant intra-specific variation in the pathogen was found for both infection success on a range of host genotypes (virulence) and components of spore production per infected host (aggressiveness) (sensu Burdon 1987). Strains did not have overall higher virulence on conspecifics of their host of origin than on strains from the heterospecific host, but they did have a significantly (c. 3 times) higher spore production per infected male host. This finding suggests that host adaptation may have evolved with respect to aggressiveness rather than virulence. The higher aggressiveness of strains on conspecifics of their host of origin resulted both from higher spore production per infected flower (spores are produced in the anthers), and greater ability to stimulate flower production on infected hosts. The latter indicates the presence of adaptive intraspecific variation in the ability of host manipulation. As transmission of the fungus is mediated by insects that are both pollinators of the host and vectors of the disease, we also assessed the effect of strains on host floral traits. Infection resulted in a reduction of inflorescence height, flower size, and nectar production per flower. Strains did not differ in their effect on nectar production, but infection with strains from S. alba resulted in a stronger reduction of inflorescence height and petal size on both host species. Vectors may therefore in principle discriminate among hosts infected by different strains and affect their efficiency of transmission. Contrary to assumptions of recent hypotheses about the role of host hybrids in the evolution of parasites, hybrids were not generally more susceptible than parental hosts. It is therefore unlikely that the rate of evolution of the pathogen on the parental species is slowed down by selection for specialization on the hybrids.     

Carabid beetle (Coleoptera: Carabidae) conservation in New Zealand

An overview of carabid beetles in the New Zealand conservation context is provided. Processes for threatened species recovery within the New Zealand Department of Conservation are outlined, and the Department’s two major beetle-related recovery documents (The Conservation Requirements for New Zealand’s Nationally Threatened Invertebrates, and The Carabid Beetle Recovery Plan) are discussed. A subjective consideration of the merits and drawbacks of both the documents, and the systems that support them, is presented, along with an option for the way forward for threatened species conservation in New Zealand.