The gastropod–crustacean connection: towards the phylogeny and evolution of the parasitic copepod family Splanchnotrophidae

Amongst the most significant metazoan taxa associated with gastropod molluscs is the endoparasitic copepod family Splanchnotrophidae. Currently it contains five genera with highly modified morphology and exclusively infesting nudibranch and sacoglossan sea slug hosts. The present study is a first approach towards reconstructing their phylogeny and evolution. Cladistic analysis of 109 morphological characters including 24 known splanchnotrophid species resulted in a fully resolved strict consensus tree that is discussed in morphological, functional, and geographical frameworks. Alternative topologies are also explored. Originating from paraphyletic Philoblennidae, the Splanchnotrophidae emerge as sister group to the genus Briarella. Unique synapomorphies, such as the bizarre body shapes and successive reduction of mouthparts, are discussed as adaptive traits to endoparasitism that evolved only once within copepods infesting shell‐less heterobranch gastropods. The ancestrally Indo‐Pacific Splanchnotrophidae split up into a clade of the still Indo‐Pacific genera Ceratosomicola and Arthurius, sister to a clade composed of the monophyletic amphi‐American genus Ismaila and European Splanchnotrophus emerging from paraphyletic Lomanoticola. Although initial radiation of Briarella and Splanchnotrophidae is likely to have involved chromodoridid nudibranch hosts, later phylogenies of parasites and their hosts are incongruent; intriguingly, host shifts from nudibranch to only distantly related sacoglossan species occurred at least two times independently. Such remarkable ecological plasticity is assumed to have driven splanchnotrophid diversification. Topological hypotheses and historical biogeographical and evolutionary scenarios inferred herein can be tested by future molecular research. © 2013 The Linnean Society of London     

Fleas associated with non‐flying small mammal communities from northern and central Chile: with new host and locality records

Fleas associated with small mammals from seven localities from northern and central Chile were assessed. We captured 352 small mammals belonging to 12 species from which we obtained 675 fleas belonging to 15 different species. The most frequently captured flea species were Neotyphloceras crassispina crassispina (n = 198) and N. chilensis (n = 175). High values of flea species richness and diversity were found in Fray Jorge National Park (NP), a north‐central Chilean site, whereas the highest values of mean abundance (MA) and prevalence were found in three diverse sites that include Los Molles River, a high altitude site located in north‐central Chile, Fray Jorge NP and Dichato, in south‐central Chile. On the other hand, high values of flea richness and diversity were found on two rodent species, Abrothrix olivacea and A. longipilis, whereas the highest values of MA and prevalence were found on Oligoryzomys longicaudatus, A. longipilis and Phyllotis xanthopygus. A total of three new host recordings, nine new localities and nine new host species and locality recordings are reported. Also, this study represents the first known record of Tetrapsyllus (Tetrapsyllus) comis in Chile and the first ecological analysis of Neotyphloceras chilensis.     

The infestation and dispersion patterns of Carcinonemertes spp. (Nemertea) on their crab hosts

Seasonal changes in the infestation and dispersion patterns of egg predatory nemerteans on their crab hosts were analyzed. Marked differences in the seasonal patterns of infestation were noted between worm species, yet, common patterns in the aggregation of the worms were found. Worm aggregation increased at the onset of the reproductive seasons of the hosts, and at the nadirs of the reproductive seasons for those hosts with year round breeding. The aggregation patterns of two worm species fluctuated with the physical environment of their estuarine hosts. Salinity changes as a result of seasonal rains may have caused changes in the underlying dispersion patterns of Carcinonemertes epialti on Hemiarapsus oregonensis, and C. mitsukurii on Portunus pelagicus. Lastly, the embryogenic cycle of the host species was significant in shaping the infestation and aggregation patterns of C. epialti on Cancer anthonyi and C. regicides on Paralithodes camtschaticus. Worm immigration and emigration were linked to crab embryogenesis and directly influenced the dispersion patterns of the worms.     

Competition,facilitation or mediation via host? Patterns of infestation of small European mammals by two taxa of haematophagous arthropods

1. We studied the effect of flea infestation on the pattern of tick (Ixodes ricinus and Ixodes trianguliceps) infestation on small mammals. 2. We asked (1) whether the probability of an individual host being infested by ticks was affected by its infestation of fleas (number of individuals and species) and (2) whether the abundance and prevalence of ticks in a host population was affected by the abundance, prevalence, level of aggregation, and species richness of fleas. 3. The probability of a host individual being infested by ticks was affected negatively by flea infestation. At the level of host populations, flea abundance and prevalence had a predominantly positive effect on tick infestation, whereas flea species richness had a negative effect on tick infestation. 4. The effect of flea infestation on tick infestation was generally greater in I. ricinus than in I. trianguliceps, but varied among host species. 5. It can be concluded that the effect of fleas on tick infestation of small mammals may be either negative or positive depending on the level of consideration and parameters involved. The results did not provide support for direct interactions between the two ectoparasite taxa, but suggested population and community dynamics and the defence system of the hosts as possible factors.     

Host infestation patterns of the massive liana Hydrangea serratifolia (Hydrangeaceae) in a Chilean temperate rainforest

As competition from lianas reduces fitness of host trees, lianas could influence community composition and structure if potential host species differ in susceptibility to infestation. We quantified infestation frequencies of Chilean temperate rainforest tree species by the massive liana Hydrangea serratifolia (H. et A.) F. Phil (Hydrangeaceae), which climbs using adhesive adventitious roots, and examined relationships with host light requirements and stem diameter. We recorded presence or absence of H. serratifolia in a random sample of 515 trees ≥10 cm diameter. Fifty‐four per cent of trees were infested by at least one individual of H. serratifolia. Although there was significant interspecific variation in infestation frequency, this variation was not systematically related to light requirements of host tree species. Probability of infestation increased with diameter for most host tree species, and old trees were found to be infested by a wide range of liana size classes, including some stems <2 cm diameter. This evidence supports the proposal that lianas which attach by adhesive roots can colonize host stems of any size.     

The influence of cactus size and previous infection on bird deposition of mistletoe seeds

Abstract Tristerix aphyllus (Loranthaceae) is a bird-dispersed mistletoe that infects several cactus species in central and north-central Chile. At Las Chinchillas National Reserve it parasitizes two species of columnar cacti (Echinopsis chilensis and Eulychnia acida) and its seeds are dispersed exclusively by the Chilean mockingbird Mimus thenca. The prevalence of T. aphyllus on cacti depended on host species and size. Echinopsis chilensis was parasitized at a higher, and E. acida at a lower, frequency than expected from their relative abundances. In both species, the incidence of T. aphyllus parasitism increased with cactus height. Birds perched and deposited seeds primarily on parasitized cacti, which resulted in an aggregated distribution of seeds. By preventing bird perching and thus reducing seed deposition, the long spines of E. chilensis and E. acida may help protect cacti against T. aphyllus infestation. However, the seeds of T. aphyllus produced a slow-growing radicle that reached up to 91 mm long in 43 days. Radicles of this length are unique among mistletoe species and may have evolved to overcome the barrier posed by cactus spines.     

<Emphasis Type="Italic">Polydora uncinata</Emphasis> (Polychaeta: Spionidae) in Chile: an accidental transportation across the Pacific

A Polydora species was found boring in shells of the abalone Haliotis discus hannai cultivated in land-based tanks in Coquimbo, Chile. Spionid polychaetes of Polydora and related genera have been reported from Chile but no worms similar to those found in abalone have been described. The abalone pest corresponds in morphology to Polydora uncinata Sato-Okoshi, 1998, a shell-boring species which was originally described from Japan and never reported from outside the country. It is suggested that occurrence of the species in Chile resulted from its accidental transportation from Japan. Adult worms were most likely transported to Coquimbo with imported abalone brood stock. Prevalence of abalone infestation by worms in Coquimbo varied substantially among cultivation tanks, reaching values as high as 98.8%. Up to 42 worms were found in one shell. The worms often caused formation of nacreous blisters which covered up to 50% of the inner shell surface. Egg capsules with developing larvae were present in female burrows. Larval development was entirely lecithotrophic, with larvae feeding on numerous nurse eggs, staying inside egg capsules until 16–17-segment stage and hatching shortly before metamorphosis. Polydora uncinata is redescribed based on individuals from Coquimbo to alert zoologists in case of accidental release of worms into Chilean coastal waters. Regardless of how the species was transported to Chile, its release to the natural ecosystem may have negative unforeseen impacts on the native fauna.     

Larval aquatic and terrestrial mites infesting a temperate assemblage of mosquitoes

We collected 22,769 adult female mosquitoes, representing 27 species, from light traps in Norfolk, Virginia (2006–2007) and examined them to assess infestation by larval mites. Mosquitoes were parasitized by two species of aquatic (Acari: Arrenuridae: Arrenurus) and three species of terrestrial mites (Acari: Erythraeidae). The prevalence of infestation varied from 0.55% (2006) to 0.17% (2007). The mean intensity of parasitism ranged from 3.6 mites per host (2006) to 1.8 mites per host (2007). The most common host species for aquatic mites was Culex erraticus, while the most common host for terrestrial mites was Anopheles quadrimaculatus. Relationships between biotic and abiotic factors were investigated in an attempt to provide insight into temporal, spatial, and interspecific variation in mite–mosquito interactions. Scanning electron microscopy was used to examine the mode of attachment for larval mites. While the prevalence of aquatic mite parasitism was correlated for Culex erraticus, the invasive mosquito, Aedes albopictus, was never parasitized through the duration of the study. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Host community structure and infestation by ixodid ticks: repeatability,dilution effect and ecological specialization

Abundance of a species in a location results from the interplay between the intrinsic properties of that species and the extrinsic properties, both biotic and abiotic, of the local habitat. Intrinsic factors promote among-population stability in abundance, whereas extrinsic factors generate variation among populations of a species. We studied (a) repeatability and (b) the effect of abundance and species richness of small mammals on the level of their infestation by larvae and nymphs of Ixodes ricinus (ecological generalist) and Ixodes trianguliceps (ecological specialist). We asked if tick infestation parameters are characteristic (=repeatable) for a particular host species or a particular stage of a particular tick species. We also asked how abundance and diversity of hosts affect the level of tick infestation on a particular host species. We predicted that the dilution effect (decrease in tick infestation levels with an increase of host abundance and/or species richness) will be better expressed in an ecological generalist, I. ricinus, than in an ecological specialist, I. trianguliceps. We found that (a) tick abundance, prevalence and aggregation were generally repeatable within tick species/stage; (b) tick abundance and prevalence, but not the aggregation level, were repeatable within host species; (c) the proportion of variance among samples explained by the differences between tick species and stages (as opposed to within-tick species and stage) was higher than that explained by the differences between host species (as opposed to within host species); and (d) the relationship between tick infestation parameters and host abundance and diversity revealed the dilution effect for I. ricinus but not for I. trianguliceps.     

The parasitism of immature stages of <Emphasis Type="Italic">Ixodes loricatus</Emphasis> (Acari: Ixodidae) on wild rodents in Argentina

Ixodes loricatus has long been considered with strict-total specificity to New World Marsupials. However, frequent findings of its immature stages on rodents suggest that these vertebrates play an important role in the tick’s life cycle. Aspects dealing with the ecology of Sigmodontinae rodents infestation by I. loricatus are unknown. To contribute to the knowledge of the ecology of this tick species, environmental factors, as well as host species, sex and age, were evaluated to find associations of immature I. loricatus infestation of the most abundant wild rodent species from riparian locations of Buenos Aires province (Argentina). A total of 290 hosts belonging to the subfamily Sigmodontinae were captured and examined for ticks at six locations. The data analysis showed that there was a preference of I. loricatus towards Akodon azarae, but only for those hosts captured in lands not prone to flooding. The results suggest that the host preference of I. loricatus is mainly related to factors such as habitat type and host density.     

Fouling community of the red king crab, <Emphasis Type="Italic">Paralithodes camtschaticus</Emphasis> (Tilesius 1815), in a subarctic fjord of the Barents sea

We examined the species composition of red king crab (Paralithodes camtschaticus) fouling communities in Dolgaya Bay, a small fjord of the Barents Sea, in August 2005 and 2006. In total, there were 13 species observed on 301 crabs collected from water depths of 5–90 m. Barnacles (Balanus crenatus; prevalence 42.9%) and blue mussels (Mytilus edulis; 11.6%) were the most common epibionts, while amphipods (Ischyrocerus commensalis) were the most common symbionts (28.6%). Infestation rates in Dolgaya Bay were different from those in an “open” area of the Barents Sea (Dalnezelenetskaya Bay), probably due to differences in hydrodynamic conditions. Differences in infestation prevalence and intensity were detected neither between male and female crabs nor between crabs collected at 5–35 m versus 90 m depths. Prevalence of common fouling species increased with host size. Amphipods I. commensalis colonized the carapace and limbs in Dolgaya Bay less frequently than in Dalnezelenetskaya Bay, probably due to interspecific competition with barnacles occupying the dorsal parts of the host. Juvenile barnacles and mussels dominated the fouling communities on the crabs. The age of barnacles did not exceed 2–4 months. However, the presence of 4-year-old mussels suggests that these older mollusks have been directly transferred from mussel beds to the hosts. Our results indicate that colonization by epibionts and symbionts is generally not disadvantageous for the crab hosts, except for some possible negative impacts of amphipods occupying the gills.     

The natural hosts for larvae and nymphs of Amblyomma neumanni and Amblyomma parvum (Acari: Ixodidae)

Based on the hypothesis that birds and rodents are important hosts for subadults of the Neotropical Amblyomma neumanni and Amblyomma parvum ticks, a survey of these type of hosts was carried out from July 2004 to March 2006, in Quilino (A. parvum) and Dean Funes (A. neumanni), Córdoba province, Argentina. Additionally, monthly tick counts were performed on cattle and goats with occasional tick search in other domestic hosts. Records of questing height of subadult ticks on vegetation were also carried out monthly. Rodents (n = 123) and birds (n = 122) captured in Dean Funes showed no infestation with A. neumanni. Apart of few nymphs found on horses, all larvae and nymphs of A. neumanni were on cattle with a larval prevalence and mean number of 22.2%, and 7.7 ± 22.52, respectively, and a prevalence of nymphs of 47.8% with a mean of 7.9 ± 18.49. The average questing height of larvae and nymphs of A. neumanni was 23.5 ± 17.1 cm and 30.7 ± 26.7 cm, respectively. A total of 138 rodents and 130 birds were captured in Quilino but the Caviidae rodent Galea musteloides carried 99.3% of larvae and 99.8% of nymphs of A. parvum, and no immature stages were detected on cattle, goat or vegetation. Tick counts on G. musteloides (n = 74) showed a prevalence of 42% and a mean number of 9.9 ± 24.83 for larvae, while nymphal infestation had a prevalence of 56.5% and a mean of 8.7 ± 11.31. Cattle appear to be suitable hosts to sustain the complete cycle of A. neumanni in nature (adult ticks infest cattle too) and questing height of subadults indicates that they are expecting to feed on medium and large-sized mammals, such as cattle and other ungulates. At least in the study site, G. musteloides is the principal host for the survival strategy of A. parvum subadults; adult ticks are common on cattle and goats. These hosts are introduced in the Neotropics but A. neumanni was able to develop a surrogate cycle independent of native hosts while A. parvum still depends on probably primeval hosts to sustain their larvae and nymphs.     

Prevalence and mean intensity of infestation by Carcinonemertes carcinophila imminuta (Nemertea: Carcinonemertidae) in the gills of Callinectes danae and Callinectes ornatus (Decapoda: Portunidae) from São Sebastião,Brazil

The nemerteans of the genus Carcinonemertes live in association with decapod crustaceans. Juveniles of Carcinonemertes carcinophila imminuta Humes, 1942 are found in the gills of their hosts. A total of 281 specimens (52 males and 229 females) of Callinectes danae Smith, 1869 and 373 (332 males and 41 females) of Callinectes ornatus Ordway, 1863 were inspected for the occurrence of Carcinonemertes carcinophila imminuta in their gills. The prevalence and mean intensity of infestation by these nemerteans with regard to sex, maturity, condition of adult female crabs (ovigerous or non ovigerous), size (carapace width) and molt stage of hosts were evaluated. A prevalence of infestation by Carcinonemertes carcinophila imminutaof 39.1% was found in the population of Callinectes danae; the mean intensity of infestation was 12.0±2.7, while intensity ranged from 1 to 268. In this host species, significant differences in prevalence were observed between males and females, juveniles and adults and ovigerous and non-ovigerous adult females. Prevalence of infestation did not differ significantly among crabs in different molt stages. Mean intensity did not differ significantly between males and females nor among crabs in different molt stages. Significant differences in mean intensity were observed between ovigerous and non-ovigerous adult females of Callinectes danae. A positive correlation was observed between prevalence of infestation and the carapace width of Callinectes danae females. The prevalence of infestation in Callinectes ornatus was 8.0%; the mean intensity of infestation was 2.7±0.4 while intensity ranged from 1 to 9. Prevalence and mean intensity of infestation did not differ significantly between males and females and juveniles and adults. Prevalence did not differ significantly among crabs in different molt stages. No correlation was observed between the prevalence of infestation and the carapace width of males and females of Callinectes ornatus.     

Physiological and ecological hosts of Popenaias popeii (Bivalvia: Unionidae): laboratory studies identify more hosts than field studies

1. Freshwater mussels are critically endangered in North America, making it important to understand their environmental requirements at all life stages. As glochidia (larvae), they attach to fish hosts where they undergo substantial mortality, making this transition important in their life cycle. Larval host fish requirements have typically been described using data from laboratory infestations to determine suitable hosts. 2. Laboratory infestations circumvent many natural barriers that prevent infestation of physiologically compatible fishes by mussel larvae. While such methods are invaluable for identifying ‘physiological hosts,’ they cannot fully describe realised ‘ecological hosts’ in the field. 3. We studied Popenaias popeii in the Black River in New Mexico, because it is of conservation concern and it is the only mussel species present, facilitating identification of glochidial infestation. To explore the difference between physiological hosts and ecological hosts, we conducted a 3‐year field study of fishes infested by P. popeii glochidia. 4. Substantially fewer fish species were infested by P. popeii in the wild (10 of 20 observed) than had been identified as physiological hosts in laboratory trials (24 of 31). We combined data on fish abundance, proportion of fish hosts infested (prevalence) and the number of glochidia per fish (intensity) and identified three fish species that probably contributed substantially more to mussel recruitment by carrying more glochidia than other host species. 5. Similarities in behaviour among these fishes allowed us to hypothesise routes of infestation, such as benthos‐feeding by catostomids, that allow glochidia to infest these hosts at higher rates than other suitable hosts. Overall, this approach provides a method of quantifying the relative importance of different species of host fish in the mussel lifecycle.     

Sex-biased parasitism,seasonality and sexual size dimorphism in desert rodents

We investigated seasonality of gender differences in the patterns of flea infestation in nine rodent species to test if sex-biased parasitism in terms of mean abundance, species richness, prevalence and the level of aggregation (a) varies among hosts and between seasons, and (b) is linked to sexual size dimorphism. Sexual size differences were significant in both summer and winter in Acomys cahirinus, Gerbillus pyramidum and Meriones crassus, and in winter only in Acomys russatus, Gerbillus dasyurus, Gerbillus nanus and Sekeetamys calurus. Sexual size dimorphism was male biased except for A. russatus in which it was female biased. Manifestation of sexual differences in flea infestation was different among hosts between seasons. A significant effect of sex on mean flea abundance was found in six hosts, on mean flea species richness in five hosts and on prevalence in two hosts. Male-biased parasitism was found in summer in one host only and in winter in five hosts. Female-biased parasitism occurred in winter in A. russatus. Gender differences in the slopes of the regressions of log-transformed variances against log-transformed mean abundances occurred in three hosts. No relationship was found between sexual size dimorphism and any parasitological parameter in any season using both conventional regressions and the method of independent contrasts. Our results suggest that sex-biased parasitism is a complicated phenomenon that involves several different mechanisms.     

The feeding process of Cimex lectularius (Linnaeus 1758) and Cimex hemipterus (Fabricius 1803) on different bloodmeal sources

The bedbugs Cimex lectularius and Cimex hemipterus are obligate hematophages in all their nymphal instars as well as in the adult stage. The efficiency with which the insects obtain blood from their hosts is directly related to their population dynamics. In the present study we compared the feeding process and salivary content in individuals of these two species when fed on different blood sources or host sites, using a cibarial pump electromyogram. Females ingested more blood than males but needed longer contact time with the host to complete the meal. The bedbug C. lectularius was more efficient than C. hemipterus in obtaining blood from mice and pigeons. With regard to the feeding site on mice, it was easier for the insects to obtain blood from the skin of the belly than that of the back. Individuals of C. hemipterus were able to maintain the cibarial pump functioning at higher frequencies for longer periods when fed on pigeons treated with anticoagulant. Although saliva from C. lectularius contained more hemeproteins and showed more anti-clotting activity its total protein content was similar to that of C. hemipterus. Overall, C. lectularius obtains a bloodmeal more efficiently from its hosts, which may have enabled this species to reach higher levels of infestation than C. hemipterus.     

Hosts and Pathogen Detection for Immature Stages of Ixodes ricinus (Acari: Ixodidae) in North-Central Spain

To determine hosts of the immature stages of a southern population of Ixodes ricinus, we trapped rodents and birds in an area of north-central Spain in May–June and August–September of 1998 and 1999. The most frequently trapped rodents were Apodemus sylvaticus (230 specimens) and Clethrionomys glareolus (99), with a larval infestation prevalence of 49% and 81%, respectively (in spring) and 21% and 41% (in summer). C. glareolus was always more heavily parasitized by larvae (mean numbers 19.8 in spring, 3.4 in summer) than A. sylvaticus (6.1 and 1.2, respectively). No nymphs were collected from the rodents. The larval distribution pattern differed considerably between rodent host species but not according to the season of the year. The most commonly trapped birds were members of the Turdidae, which had infestation incidences of about 70% (larvae) and 10% (nymphs) in spring and 20–15% in summer. We also used molecular methods for the identification of reservoir hosts of the larval ticks and of the pathogens they contained, from nymphs collected in the same site in June of 2002 and 2003. The study showed that most of the ticks had fed on birds and few on rodents and ruminants. Pathogens detected in these ticks included Babesia microti, Borrelia garinii, B. valaisiana and B. afzelii. Borrelia afzelii was detected in a tick that had apparently fed on a wild boar.     

Parasitism by Pteromalus cerealellae (Hymenoptera: Pteromalidae) on the Cowpea weevil, Callosbruchus maculatus (Coleoptera: Bruchidae): Host density, temperature effects, and host finding ability

Studies on interactions between a larval parasitoid, Pteromalus cerealellae (Boucek) and one of its hosts, Callosobruchus maculatus (F.) were carried out in the laboratory. The number of host larvae parasitized by P. cerealellae increased with host larvae at low densities and tended to a plateau at a density of 25 larvae per female parasitoid. Each parasitoid was able to parasitize more hosts and produced more offspring at 20 and 25 °C than at 30 °C. The number of non-infested seeds mixed with seeds infested with the last instar of C. maculatus did not preclude P. cerealellae from identifying infested seeds and attacking larvae inside them. When infested seeds were tightly packed, several host larvae escaped parasitism. P. cerealellae may be a useful biological control agent in newly harvested cowpea with low C. maculatus infestation, and lowering the temperature of the storage system may enhance the effectiveness of this parasitoid.     

Occurrence and performance of the aspen blotch miner,Phyllonorycter salicifoliella,on three host-tree species

Summary Larvae of the aspen blotch miner, Phyllonorycter salicifoliella Chambers (Lepidoptera: Gracillariidae), feed within leaves of three host-tree species in north-central Minnesota, USA. Far more individuals occur on Populus tremuloides than on P. balsamifera or P. grandidentata. We tested whether this pattern of host use reflected variable performance among alternative hosts by examining survivorship, sources of mortality, pupal mass, feeding efficiency, and development time of miners on each tree species. We also determined foliar water, nitrogen, condensed tannin, and phenolic glycoside content of host trees to test if host-tree chemical attributes were responsible for differences in performance. There was no significant difference in egg-to-adult survival among miners on different hosts, although dominant sources of mortality did vary. Miners on P. grandidentata suffered less parasitism and more predation than those on the other hosts, even though most parasitoid species attacked miners on all hosts. The other performance parameters varied among host species, but not in a consistent pattern. Pupal mass was greatest on P. tremuloides and P. balsamifera, the hosts with comparatively high foliar nitrogen and low phenolic glycoside concentrations. However, feeding efficiency was greatest and development time shortest for miners on P. grandidentata. Thus, pupal mass was the only index of performance maximized on P. tremuloides, the most commonly used host. Infrequent occurrence of Phyllonorycter salicifoliella on P. grandidentata results in part from phenological differences between this and the other host species. Low oviposition rates on P. balsamifera are correlated with low abundance of this host at the study site and a phenolic glycoside profile different from that of the other host species.