Determinants of helminth aggregation in natural host populations: individual differences or spatial heterogeneity?

We assessed the importance of spatial heterogeneity for the aggregation of helminth populations in the bank vole Clethrionomys glareolus (Arvicolinae) using a previously published method which allows to analyse parasite aggregation at two host population levels, i.e., within and between spatial samples of the host population.
In the main empirical material from Finnish Lapland (Pallasjärvi), all five helminth species were significantly aggregated within study sites, but only three rare species showed significant aggregation among sites. In all helminth species, the total aggregation was primarily (79-98%) determined by aggregation within sites, i.e., between host individuals. Despite the larger spatial scale and more heterogeneous landscape, the comparative material from South Finland (Luhanka) supported the generally high proportion of the total aggregation due to within-site heterogeneity.
Despite the clear interspecific differences in patchiness of helminth populations, the proportion of the total aggregation due to among-site heterogeneity did not vary significantly among helminth species. The results indicate a link between aggregation at two population levels; species that showed strong within-site aggregation were also characterised by pronounced spatial heterogeneity and significant among-site aggregation.     

Diet divergence in two sympatric congeneric butterflies: Community or species level phenomenon?

Summary Two species ofEuphydryas butterflies were studied in California, USA, and showed considerable diet overlap at the species level. They utilize many of the same plant genera for oviposition. However,E. editha is less likely to use woody perennials than isE. chalcedona.Both butterfly species are known to specialize on different host plants in different populations, so species level divergence may not be a good predictor of community level divergence. Within five communities,E. editha andE. chalcedona showed no dietary overlap. A major component of the niche ofE. editha in one community was occupied byE. chalcedona in a second community, even though both butterfly species occupied both communities. These resource use patterns indicate that community level interactions may affect diet divergence. The degree to which divergence within communities is greater (or less) than expected from a species level comparison may be used to provide a measure of community organization. Equations are given in the Appendix for calculating overlap probabilities from presence/absence types of data; in this study, presence is oviposition on a particular plant species, absence is no oviposition on that plant species. Given the various assumptions of the model,E. editha andE. chalcedona show significant community level components of their dietary divergence in the areas studied. However, in some other communitiesE. editha andE. chalcedona do share host plant species. Therefore, we could not demonstrate community level divergence conclusively, nor has this been demonstrated for any other pair of insect herbivore species. We do not know whether this is because the phenomenon is truly rare or just very hard to demonstrate.     

Mud-puddling behavior in tropical butterflies: in search of proteins or minerals?

We experimentally investigated the attraction of adult butterflies to moist soil and dirt places (a behavior termed `mud-puddling') in two species-rich tropical communities on the island of Borneo. At a rain forest site, 227 individuals (46 species) were attracted to the baits, compared to 534 individuals (54 species) at a farmland site. With one single exception, all attracted butterflies were males. Of various salt and amino acid solutions, only sodium was accepted, but overall, albumin solutions turned out to be the most attractive puddling resource. Butterfly families differed consistently in their resource preferences. Representatives of the families Papilionidae and Pieridae more often visited NaCl solutions, but still accepted albumin, whereas representatives of the Nymphalidae, Hesperiidae and, in particular, Lycaenidae preferred the protein resource. In experiments using decoys prepared from pinned butterfly specimens, representatives of the Papilionidae and Pieridae were more strongly attracted to baits provided with decoys made from conspicuous, medium-sized yellow Eurema species (Pieridae), whereas dummies made from small, cryptically colored lycaenids (Prosotas and Caleta species) were ineffective. Decoys did not influence the attraction of lycaenid butterflies towards baits. Hence, visual cues play an important role in locating puddling resources for papilionids and pierids, while for lycaenid butterflies searching for nitrogen sources, olfactory cues emitted by decaying organic matter are more likely to be important. The strong attraction of male butterflies to nitrogen-rich resources suggests that, as in the case of sodium, these nutrients may increase reproductive success. Received: 5 October 1998 / Accepted: 7 December 1998     

Evolution of host specificity in fleas: is it directional and irreversible?

Evolutionary trends in the evolution of host specificity have been the focus of much discussion but little rigorous empirical testing. On the one hand, specialization is often presumed to lead irreversibly into evolutionary dead ends and little diversification; this would mean that generalists might evolve into specialists, but not vice versa. On the other hand, low host specificity may limit the risk of extinction and provide more immediate fitness benefits to parasites, such that selection may favour evolution toward a generalist strategy. Here, we test for directionality in the evolution of host specificity using a large data set and phylogenetic information on 297 species of fleas parasitic on small mammals. The analyses determined whether host specificity, measured both as the number of host species exploited and their taxonomic diversity, was related to clade rank of the flea species, or the number of branching events between an extant species and the root of the phylogenetic tree (i.e., the total path length from the root of the tree to the species). Based on regression analyses, we found positive relationships between the number of host species used and clade rank across all 297 species, as well as within one (Hystrichopsyllidae) of four large families and one of seven large genera investigated separately; in addition, we found a positive relationship between the taxonomic diversity of host species used and clade rank in another of the seven genera. These results suggest a slight evolutionary trend of decreasing host specificity. Using a much more conservative likelihood ratio test, however, a random walk, or null model, of evolution could not be discarded in favour of the directional trends in all cases mentioned above. Still, these results suggest that host specificity may have tended to decrease in many flea lineages, a process that could have been driven by the benefits of exploiting a wide range of host species.     

Do major host shifts spark diversification in butterflies?

The Escape and Radiate Hypothesis posits that herbivorous insects and their host plants diversify through antagonistic coevolutionary adaptive radiation. For more than 50 years, it has inspired predictions about herbivorous insect macro‐evolution, but only recently have the resources begun to fall into place for rigorous testing of those predictions. Here, with comparative phylogenetic analyses of nymphalid butterflies, we test two of these predictions: that major host switches tend to increase species diversification and that such increases will be proportional to the scope of ecological opportunity afforded by a particular novel host association. We find that by and large the effect of major host‐use changes on butterfly diversity is the opposite of what was predicted; although it appears that the evolution of a few novel host associations can cause short‐term bursts of speciation, in general, major changes in host use tend to be linked to significant long‐term decreases in butterfly species richness.     

How do two specialist butterflies determine growth and biomass of a shared host plant?

Although insect herbivory can modify subsequent quantity and quality of their host plants, change in plant quantity following herbivory has received less attention than plant quality. In particular, little is known about how previous herbivore damage determines plant growth and biomass in an insect species-specific manner. We explored whether herbivore species-specific food demand influences plant growth and biomass. To do this, we conducted a series of experiments and field survey using two specialist butterflies, Sericinus montela and Atrophaneura alcinous, and their host plant, Aristolochia debilis. It is known that A. alcinous larva requires four times more food resources to fulfill its development than S. montela larva. Despite that A. alcinous larvae imposed greater damage on plants than S. montela larvae, plant growth did not differ due to herbivory by these species both in single and multiple herbivory events. On the other hand, total aboveground biomass of the plants was reduced more by A. alcinous than S. montela feeding regardless of the number of herbivory events. Feeding on plants with a history of previous herbivory neither decreased nor increased larval growth. Our results suggest that food demand of the two butterfly species determined subsequent plant biomass, although the plant response may depend on tolerance of the host plant (i.e., ability to compensate for herbivore damage). Such difference in the effects of different herbivore species on host plant biomass is more likely to occur than previously thought, because food demand differs in most herbivore species sharing a host plant.     

Ability to gall: the ultimate basis of host specificity in fig wasps?

1. Fig trees (Ficus spp.) and their host‐specific pollinator fig wasps (Agaonidae) are partners in an obligate mutualism. Receptive phase figs release specific volatiles to attract their pollinators, and this is generally effective in preventing pollinator species from entering figs of the wrong hosts. 2. If entry is attempted into atypical host figs, then ostiole size and shape and style length may also prevent reproduction. In spite of these barriers, there is increasing evidence that fig wasps enter atypical hosts, and that this can result in hybrid seed and fig wasp offspring. 3. This study examines the basis of pollinator specificity in two dioecious fig species from different geographical areas. Kradibia tentacularis pollinates Ficus montana in Asia. Ficus asperifolia from East Africa is closely related but is pollinated by a different species of Kradibia. 4. In glasshouses, K. tentacularis was attracted to its normal host, F1s and backcrosses, but only rarely entered figs of F. asperifolia. Foundresses were able to lay eggs in hybrids, backcrosses, and F. asperifolia, although flower occupancy was lowest in F. asperifolia figs and intermediate in hybrids. 5. The fig wasp failed to reproduce in female F. montana, male F. asperifolia, and male F1s, and most but not all backcrosses to F. montana. This was a result of the failure to initiate gall production. 6. Host specificity in this fig wasp is strongly influenced by host volatiles, but the ability to gall may be the ultimate determinant of whether it can reproduce.     

Sexual differences in habitat use by small mammals: evolutionary strategy or reproductive constraint?

Summary This study examines the hypothesis that intersexual differences in habitat use by small mammals is an adaptive strategy. Specifically, I evaluate the occurrence of sexual differences in habitat by Peromyscus and Microtus, and test possible adaptive mechanisms which may lead to such differences.White-footed mice and meadow voles were live-trapped and microhabitat quantified in four habitats. Sexual differences in microhabitat use were found in three of 12 comparisons. These differences may be related to density dependent resource subdivision and habitat heterogencity, but neither of these can account completely for the observed patterns. Sexual differences in habitat do not appear to be an evolutionary strategy maximizing reproductive effort by females, or reducing predation. These sexual differences may reduce intraspecific resource overlap, or may simply reflect reproductive constraints limiting female habitat use to suitable nesting areas.     

Genetic structure and gene flow in French populations of two Ostrinia taxa: host races or sibling species?

Most models of ecological speciation concern phytophagous insects in which speciation is thought to be driven by host shifts and subsequent adaptations of populations. Despite the ever-increasing number of studies, the current evolutionary status of most models remains incompletely resolved, as estimates of gene flow between taxa remain extremely rare. We studied the population genetics of two taxa of the Ostrinia genus--one feeding mainly on maize and the other on mugwort and hop--occurring in sympatry throughout France. The actual level of divergence of these taxa was unknown because the genetic structure of populations had been investigated over a limited geographical area and the magnitude of gene flow between populations had not been estimated. We used 11 microsatellite markers to investigate the genetic structure of populations throughout France and the extent of gene flow between the two Ostrinia taxa at several sites at which they are sympatric. We observed clear genetic differentiation between most populations collected on the typical respective hosts of each taxon. However, populations displaying intermediate allelic frequencies were found on hop plants in southern France. Individual assignments revealed that this result could be accounted for by the presence of both taxa on the same host. Gene flow, estimated by determining the proportion of hybrids detected, was low: probably<1% per generation, regardless of site. This indicates that the two Ostrinia taxa have reached a high level of genetic divergence and should be considered sibling species rather than host races.     

Oviposition specificity in single vs. cluster egg-laying butterflies: a discrimination phase in Colias eurytheme?

Summary Oviposition preferences and rates of the pierid butterfly Colias eurytheme were examined in laboratory tests. Oviposition rates with only a non-preferred host plant present were not lower than rates observed when both preferred and non-preferred hosts were present. There was no correlation between strength of preference in simultaneous choice tests and the extent to which oviposition rate declined when the preferred host was absent. The results are consistent with the hypothesis that butterflies which lay their eggs singly will accept non-preferred hosts for oviposition more readily than cluster egg-layers.     

Increased host acceptance in experienced females of the parasitoid Brachymeria intermedia: which types of oviposition behaviour contribute to experience?

Brachymeria intermedia (Nees) (Hymenoptera: Chalcididae) is a solitary endoparasitoid of the gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae). The probability that a wasp will parasitise this host is increased if she had a previous oviposition experience. We investigated which types of pre-oviposition and oviposition behaviour induced this change by comparing the acceptance behaviour of wasps with zero to five oviposition experiences as well as wasps with various partial host handling experiences (from no contact with a host to one complete oviposition sequence). The percentage of females accepting the host increased gradually with both the number of previous oviposition experiences and the amount of handling of a single host. Furthermore, the naive females were less likely (44%) to walk to the host than females with a variety of experiences (72–100%). A single antennal contact with the host was sufficient to increase the probability of walking to the host. Additional handling and additional oviposition experiences further elevated the propensity to oviposit. Thus, the modification of the acceptance behaviour through experience was essentially a gradual process in which antennal contact with a pupa was a major element. It is suggested that naive wasps learned the host odour when they first antennated a pupa. As a result, they were attracted to the host odour in subsequent encounters.     

Does host plant quality affect the oviposition decisions of an omnivore?

Optimal oviposition theory predicts a positive relationship between female preference for oviposition hosts and offspring performance. Interspecies effects on oviposition preference have been widely investigated, especially for herbivores. However, intraspecies variation, such as nitrogen content, might also influence female preference for oviposition hosts and subsequent offspring performance. To evaluate this possibility, we investigated the oviposition preference of a zoophytophagous omnivore and the development and survival of its nymphs on a single species of host plant that varied in nitrogen content. In choice and no‐choice experiments without prey, female omnivores were allowed to oviposit on plants that had been fertilized using 4 rates of nitrogen fertilizer (39, 78, 156, and 311 mg/L nitrogen) for 72 h. After 72 h, the most females were found on tomato plants receiving high concentrations of nitrogen fertilizer and more eggs were laid on those plants. First instar nymphs developed more rapidly on high‐nitrogen plants and third instar nymphs developed faster on low‐nitrogen plants. Plant nitrogen did not affect nymph survival to the adult stage, or the probability of survival over time. Although female omnivores did discriminate between potential oviposition hosts based on plant nitrogen, their choices did not significantly impact nymph development or survival. This is the first study to show that intraspecies variation in nitrogen content between plants affects the oviposition preference of female omnivores, but not offspring performance.     

Non-target plant use by a weed biocontrol agent in idaho: host expansion or opportunistic behavior?

Larinus curtus Hochhuth (Coleoptera: Curculionidae) was first introduced into the western United States from Greece for the biological control of yellow starthistle (YST), Centaurea solstitialis L., in 1992. The discovery of L. curtus adults in the open flowerheads of safflower (SF), Carthamus tinctorius L., near Lewiston, Idaho in 2007 suggested this weevil might be expanding its host range to include a non-target crop species closely related to YST. In 2008 field plots near the 2007 observation site, 92 L. curtus adults fed in open SF flowerheads (pollen feeding and minor feeding on corolla tubes). No eggs were found in the ovarioles of 19 pollen-feeding females. No eggs, larvae, or evidence of larval feeding were detected in 39 tagged SF capitula, and no adults emerged from approximately 7,135 post-flowering SF capitula. These collective results are not indicative of an expanding developmental host-range of L. curtus. Also, they are consistent with pre-release host-specificity test results.     

US specificity of occasion setting: Hierarchical or configural learning?

Four experiments in rats examined whether occasion setters and target CSs play qualitatively different roles in occasion-setting discriminations. Two visual occasion setters, A and B, signalled reinforcement of two auditory target CSs, x and y, with sucrose and oil (A…x→suc, B…y→oil, A-, B-, x-, y-); in addition two transfer CSs w and z were paired with sucrose and oil (w→suc, z→oil). When w and z were substituted for x and y (A…w, B…w, A…z, B…z) more responding was observed when both stimuli had been paired with the same outcome (Experiments 1 and 3a). No effect was observed when two visual \"pseudo-occasion setters\", C and D (paired with sucrose and oil in a trace relation to the US:C…→suc, D…→oil), were substituted for the occasion setters A and B (C…x, D…x, C…y, D…y; Experiments 2, 3b and 4). These results could not be explained in terms of Pavlovian summation: responding to combinations of Pavlovian CSs paired with same or different outcomes was either the same, or lower when both stimuli had been paired with the same outcome (Experiment 4). Implications of these results for theories of occasion setting and configural learning are discussed.     

Does previous experience enhance foraging on a particular host in a polyphagous frugivore?

Prior oviposition or feeding experience with a host plant has been proposed to improve foraging efficiency for polyphagous insect herbivores. Many laboratory-based experiments have attributed certain modified behavioural responses, such as an improved ability to locate a host, as evidence for the benefits of learning. However, few studies have considered the negative ramifications of learning, particularly under field conditions rather than laboratory conditions. In the current study, using the polyphagous fruit fly Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), we explore both the positive and negative consequences of modified host selection resulting from prior oviposition experience. In field cage experiments, prior oviposition experience on a given host increased the selection of that host despite its abundance and its suitability for offspring development. For example, in a field cage that contained significantly more blueberry (poor larval host) than guava (good larval host), prior experience on guava resulted in more flies selecting and accepting guava than did naïve flies (a presumed positive foraging outcome). However, the opposite also held, that prior experience on blueberry led to increased blueberry use, even when guava was more abundant. In unrestrained flies, prior experience dramatically improved recapture rate with flies locating the training fruit. The results highlight the potential benefits of learning, but also identify that learning can be a disadvantage if prior experience on a poor host leads to flies repeatedly using that host over a more suitable host.     

Evolutionary implications of host–pathogen specificity: the fitness consequences of host life history traits

Pathogens and parasites can be strong agents of selection, and often exhibit some degree of genetic specificity for individual host strains. Here we show that this host–pathogen specificity can affect the evolution of host life history traits. All else equal, evolution should select for genes that increase individuals' reproduction rates or lifespans (and thus total reproduction per individual). Using a simple host–pathogen model, we show that when the genetic specificity of pathogen infection is low, host strains with higher reproduction rates or longer lifespans drive slower-reproducing or shorter-lived host strains to extinction, as one would expect. However, when pathogens exhibit specificity for host strains with different life history traits, the evolutionary advantages of these traits can be greatly diminished by pathogen-mediated selection. Given sufficient host–pathogen specificity, pathogen-mediated selection can maintain polymorphism in host traits that are correlated with pathogen resistance traits, despite large intrinsic fitness differences among host strains. These results have two important implications. First, selection on host life history traits will be weaker than expected, whenever host fitness is significantly affected by genotype-specific pathogen attack. Second, where polymorphism in host traits is maintained by pathogen-mediated selection, preserving the genetic diversity of host species may require preserving their pathogens as well. This revised version was published online in November 2006 with corrections to the Cover Date.