Isolation of Psoroptes scab mite microsatellite markers (Acari: Psoroptidae)

Nine microsatellite markers have been isolated from the scab mite, Psoroptes ovis. These markers have been tested for polymorphism in individual Psoroptes mites originating from two hosts, the European rabbit, Oryctolagus cuniculus, and sheep, Ovis aries. No definitive picture of Psoroptes species’ status or interrelationships exists. This study provides the basis for a new molecular system to elucidate the systematics of groupings within the genus Psoroptes, allowing us to clarify the population dynamics and epidemiology of the mites causing sheep scab world wide.     

Species of the genus Psoroptes (Acari: Psoroptidae): A taxonomic consideration

The biosystematic status of mite species belonging to the genus Psoroptes Gervais, 1841 is difficult to determine by phenotypic methods and has been subject to taxonomic revisions and ongoing debate. At present, the existence of five species, P. cuniculi (Delafond, 1859), P. ovis (Hering, 1838), P. equi (Hering, 1838), P. cervinus Ward, 1915 and P. natalensis Hirst, 1919, is generally accepted. This classification is based mainly on the host species, the localization of the mites on their hosts and morphological characters of male mites. However, a critical review of the literature indicates that the features used to discriminate between the five species are not unequivocal: (a) the localization of mite populations on host animals is not completely strict, (b) the lengths of the outer opisthosomal setae of male mites, which are the main morphological features used for species discrimination, overlap between the five postulated species, and (c) host specificity cannot be deduced from results of transfer experiments. Rather, conspecificity of the members of the genus Psoroptes has to be presumed which is supported by molecular genetic analyses. On these grounds and on rules of priority P. cervinus Ward, 1915, P. cuniculi (Delafond, 1859), P. natalensis Hirst, 1919 and P. ovis (Hering, 1838) are seen as synonyms of P. equi (Hering, 1838).     

Sex-biased parasitism is not universal: evidence from rodent–flea associations from three biomes

The distribution of parasites among individual hosts is characterised by high variability that is believed to be a result of variations in host traits. To find general patterns of host traits affecting parasite abundance, we studied flea infestation of nine rodent species from three different biomes (temperate zone of central Europe, desert of Middle East and tropics of East Africa). We tested for independent and interactive effects of host sex and body mass on the number of fleas harboured by an individual host while accounting for spatial clustering of host and parasite sampling and temporal variation. We found no consistent patterns of the effect of host sex and body mass on flea abundance either among species within a biome or among biomes. We found evidence for sex-biased flea infestation in just five host species (Apodemus agrarius, Myodes glareolus, Microtus arvalis, Gerbillus andersoni, Mastomys natalensis). In six rodent species, we found an effect of body mass on flea abundance (all species mentioned above and Meriones crassus). This effect was positive in five species and negative in one species (Microtus arvalis). In M. glareolus, G. andersoni, M. natalensis, and M. arvalis, the relationship between body mass and flea abundance was mediated by host sex. This was manifested in steeper change in flea abundance with increasing body mass in male than female individuals (M. glareolus, G. andersoni, M. natalensis), whereas the opposite pattern was found in M. arvalis. Our findings suggest that sex and body mass are common determinants of parasite infestation in mammalian hosts, but neither of them follows universal rules. This implies that the effect of host individual characteristics on mechanisms responsible for flea acquisition may be manifested differently in different host species.     

Serodiagnostic antibody responses to Psoroptes sp. infestations in bighorn sheep

The antibody responses of bighorn sheep (Ovis canadensis) infected with Psoroptes sp. mites were investigated by enzyme linked immunosorbent assay on western blots of P. cuniculi antigens. Serum from 20 Psoroptes sp.-infested bighorn sheep (O. canadensis mexicana, O. canadensis nelsoni, O. canadensis canadensis) from New Mexico, Nevada, California, and Idaho reacted strongly with mite antigens ranging from 12 to 34 kd. Serum from 35 Psoroptes sp.-free bighorn sheep of unknown tick infestation status and from three Psoroptes sp.-free bighorn sheep infested with Dermacentor hunteri ticks did not react with these antigens. Psoroptes sp.-specific antibody responses were present throughout a 16 mo period in one infected bighorn sheep, but were not detectable 8 mo following successful treatment. These results demonstrate that specific serodiagnosis of Psoroptes sp. infestation is feasible in bighorn sheep and suggest that antibody responses are indicative of current or recent infestation.     

On the Front Line: Quantitative Virus Dynamics in Honeybee (Apis mellifera L.) Colonies along a New Expansion Front of the Parasite Varroa destructor

Over the past fifty years, annual honeybee (Apis mellifera) colony losses have been steadily increasing worldwide. These losses have occurred in parallel with the global spread of the honeybee parasite Varroa destructor. Indeed, Varroa mite infestations are considered to be a key explanatory factor for the widespread increase in annual honeybee colony mortality. The host-parasite relationship between honeybees and Varroa is complicated by the mite''s close association with a range of honeybee viral pathogens. The 10-year history of the expanding front of Varroa infestation in New Zealand offered a rare opportunity to assess the dynamic quantitative and qualitative changes in honeybee viral landscapes in response to the arrival, spread and level of Varroa infestation. We studied the impact of de novo infestation of bee colonies by Varroa on the prevalence and titres of seven well-characterised honeybee viruses in both bees and mites, using a large-scale molecular ecology approach. We also examined the effect of the number of years since Varroa arrival on honeybee and mite viral titres. The dynamic shifts in the viral titres of black queen cell virus and Kashmir bee virus mirrored the patterns of change in Varroa infestation rates along the Varroa expansion front. The deformed wing virus (DWV) titres in bees continued to increase with Varroa infestation history, despite dropping infestation rates, which could be linked to increasing DWV titres in the mites. This suggests that the DWV titres in mites, perhaps boosted by virus replication, may be a major factor in maintaining the DWV epidemic after initial establishment. Both positive and negative associations were identified for several pairs of viruses, in response to the arrival of Varroa. These findings provide important new insights into the role of the parasitic mite Varroa destructor in influencing the viral landscape that affects honeybee colonies.     

Differential Water Mite Parasitism,Phenoloxidase Activity,and Resistance to Mites Are Unrelated across Pairs of Related Damselfly Species

Related host species often demonstrate differences in prevalence and/or intensity of infection by particular parasite species, as well as different levels of resistance to those parasites. The mechanisms underlying this interspecific variation in parasitism and resistance expression are not well understood. Surprisingly, few researchers have assessed relations between actual levels of parasitism and resistance to parasites seen in nature across multiple host species. The main goal of this study was to determine whether interspecific variation in resistance against ectoparasitic larval water mites either was predictive of interspecific variation in parasitism for ten closely related species of damselflies (grouped into five “species pairs”), or was predicted by interspecific variation in a commonly used measure of innate immunity (total Phenoloxidase or potential PO activity). Two of five species pairs had interspecific differences in proportions of individuals resisting larval Arrenurus water mites, only one of five species pairs had species differences in prevalence of larval Arrenurus water mites, and another two of five species pairs showed species differences in mean PO activity. Within the two species pairs where species differed in proportion of individuals resisting mites the species with the higher proportion did not have correspondingly higher PO activity levels. Furthermore, the proportion of individuals resisting mites mirrored prevalence of parasitism in only one species pair. There was no interspecific variation in median intensity of mite infestation within any species pair. We conclude that a species’ relative ability to resist particular parasites does not explain interspecific variation in parasitism within species pairs and that neither resistance nor parasitism is reflected by interspecific variation in total PO or potential PO activity.     

Reproduction of ectoparasitic mites in a coevolved system: Varroa spp.—Eastern honey bees,Apis cerana

Parasite host shifts can impose a high selective pressure on novel hosts. Even though the coevolved systems can reveal fundamental aspects of host–parasite interactions, research often focuses on the new host–parasite relationships. This holds true for two ectoparasitic mite species, Varroa destructor and Varroa jacobsonii, which have shifted hosts from Eastern honey bees, Apis cerana, to Western honey bees, Apis mellifera, generating colony losses of these pollinators globally. Here, we study infestation rates and reproduction of V. destructor and V. jacobsonii haplotypes in 185 A. cerana colonies of six populations in China and Thailand to investigate how coevolution shaped these features. Reproductive success was mostly similar and low, indicating constraints imposed by hosts and/or mite physiology. Infestation rates varied between mite haplotypes, suggesting distinct local co‐evolutionary scenarios. The differences in infestation rates and reproductive output between haplotypes did not correlate with the virulence of the respective host‐shifted lineages suggesting distinct selection scenarios in novel and original host. The occasional worker brood infestation was significantly lower than that of drone brood, except for the V. destructor haplotype (Korea) from which the invasive lineage derived. Whether mites infesting and reproducing in atypical intraspecific hosts (i.e., workers and queens) actually predisposes for and may govern the impact of host shifts on novel hosts should be determined by identifying the underlying mechanisms. In general, the apparent gaps in our knowledge of this coevolved system need to be further addressed to foster the adequate protection of wild and managed honey bees from these mites globally.     

EFFECT OF ECTOPARASITIC MITES ON SEXUAL SELECTION IN A SONORAN DESERT FRUIT FLY

We conducted a field study and a laboratory experiment to test whether ectoparasitic mites, Macrocheles subbadius, generate parasite-mediated sexual selection in the Sonoran desert endemic fruit fly, Drosophila nigrospiracula. Male flies gather on the outer surfaces of necrotic saguaro cacti where they engage in male–male competitive interactions and vigorous female-directed courtship. At these sites, operational sex ratios were significantly skewed toward males. The degree to which mites were aggregated among flies varied across the 25 fly populations sampled. The degree of mite aggregation across fly populations was strongly positively related to the mean number of mites per fly (intensity of infestation). Both the intensity and prevalence of infestation (fraction of flies infested) increased with the age of the cactus necrosis. Infested flies of both sexes were significantly less likely to be found in copula than uninfested flies, and mean intensity of infestation was significantly more pronounced in noncopulating than in copulating flies. The effect of attached mites on copulatory success exhibited dose-dependency, and this effect was more stringent in males: males or females with more than two and four mites, respectively, were never found in copula. The magnitude of parasite-mediated sexual selection was estimated for 12 fly populations by calculating selection differentials for each sex separately. The relation between intensity of infestation and magnitude of parasite-mediated sexual selection was stronger in males but significant for both sexes. We also assayed copulatory success of field-caught males in the laboratory, both during infestation and after experimental removal of mites. Males infested with two mites copulated less frequently than uninfested individuals, and in mating trials after mites had been removed, previously infested males copulated as many times as flies with no history of infestation. These findings, and the lack of difference in the number of mite-induced scars on copulating and single individuals in nature, strongly suggest that the reduced copulatory success of infested flies is attributable to an effect of mites per se, rather than to a character correlated with parasitism or previous parasite infestation.     

Morphological and molecular comparison of host-derived populations of parasitic Psoroptes mites

Infestation by parasitic Psoroptes mites (Acari: Psoroptidae) is an important cause of economic loss and welfare problems in livestock in many areas of the world. At least five species within this genus have been recognized, based on the host infested, the infestation site and differences in length of the opisthosomal setae of adult male mites. Here the integrity of these species is considered by subjecting populations of mites from a range of host species and geographical locations to simultaneous morphological and molecular genetic analyses. Morphological analysis showed that there were significant differences in shape and size between mite populations from different hosts, and that length of the outer opisthosomal setae in males and the homologous seta in females were the most important distinguishing character in adults. However, considerable variation in outer opisthosomal seta length was evident within and between populations of mites, and differences were not clearly related to host-species or geographical origin and did not support the accepted species differences. Molecular characterization using sequence data from the mitochondrial second internal transcribed spacer (ITS-2) region and microsatellite markers found little or no consistent host-related variation between the mite population samples. The results suggest that there is no case for considering the Psoroptes mites from the different hosts examined as separate species and that the morphological variation observed therefore may represent phenotypic adaptation to the local microenvironment on particular species of host.     

Occurrence and seasonal prevalence of the coconut mite, Aceria guerreronis (Eriophyidae), and associated arthropods in Oman

The coconut palm is an important crop in the sub arid coastal plain of Dhofar, Oman, for the high demand for its nut water and its use as ornamental plant. Damage of coconut fruits by the eriophyid mite Aceria guerreronis Keifer was first reported in that region in the late 1980s, but background information about the ecology of the pest in Oman was missing. Four surveys were conducted in different seasons from 2008 to 2009, to assess the distribution and prevalence of the coconut mite and its damage as well as the presence of natural enemies. Infestation by the coconut mite was conspicuous on most (99.7 %) palm trees, with 82.5 % damaged fruits. The average (±SE) density of coconut mites per fruit was 750 ± 56; this level of infestation led to the incidence of over 25 % of surface damage on more than half of the fruits. The mite appeared more abundant at the end of the cold season through the summer. No significant differences were observed between infestation levels on local varieties, hybrids and on dwarf varieties. Neoseiulus paspalivorus (De Leon), Cydnoseius negevi (Swirski & Amitai) and Amblyseius largoensis (Muma) were the predatory mites found under the bracts of over 30 % of the coconut fruits and on 68 % of the coconut trees. Considering all sampling dates and all varieties together, average (± SE) phytoseiid density was 1.4 ± 1.19 per fruit. Other mites found in the same habitat as A. guerreronis included the tarsonemids Steneotarsonemus furcatus De Leon and Nasutitarsonemus omani Lofego & Moraes. The pathogenic fungus Hirsutella thompsonii Fisher was rarely found infecting the coconut mite in Dhofar. Other fungal pathogens, namely Cordyceps sp. and Simplicillium sp., were more prevalent.     

Mite pests and their predators on seven vegetable crops (Arachnida: Acari)

During this study the frequency of occurrence and dominance of phytophagous and predatory mites harboring seven vegetable crops in Egypt, namely common bean, cowpea, eggplant, okra, squash, sweet pepper and sweet potato during 2017–2018 were investigated to identify predatory mites that might be useful for the biological control of the phytophagous mites. Three phytophagous and nine predatory mite species were surveyed. The two spotted spider mite Tetranychus urticae Koch of the family Tetranychidae was the dominant pest on these vegetables, while phytoseiids Phytoseiulus persimilis (Athias- Henriot), Typhlodromips swirskii (Athias- Henriot) and Euseius scutalis Chant were the dominant predators. The population of the native or indigenous phytoseiid mite fauna in Egypt such as Phytoseiulus persimilis could be considered as a good biocontrol agent and a part of the Integrated Pest Management (IPM) program in the future. Mite fauna of Egypt especially local populations of Phytoseiulus persimilis can be considered for implementation in future Integrated Pest Management (IPM).     

Analysis of ectoparasites (chigger mites,gamasid mites,fleas and sucking lice) of the Yunnan red‐backed vole (Eothenomys miletus) sampled throughout its range in southwest China

The Yunnan red‐backed vole Eothenomys miletus (Rodentia: Cricetidae) is an endemic rodent species and reservoir host of zoonoses in southwest China. Based on a large host sample (2463 voles collected from 39 localities between 2001 and 2013), a general analysis of four categories of ectoparasite (fleas, sucking lice, chigger mites and gamasid mites) on E. miletus across its entire range of distribution was made. This analysis identified a total of 71 895 ectoparasites belonging to 320 species (30 species of flea, 9 of sucking louse, 106 of gamasid mite and 175 of chigger mite) with a high prevalence (87%), mean abundance (29.19) and mean intensity (33.69). Of the 18 vector species of zoonoses found on E. miletus, the flea Ctenophthalmus quadratus (Siphonaptera: Hystrichopsyllidae) and chigger mite Leptotrombidium scutellare (Trombidiformes: Trombiculidae) were the dominant species; these are the main vectors of zoonoses in China. All of the dominant parasite species showed an aggregated distribution pattern. Male voles harboured more species of parasite than females. Chigger mites represented the most abundant species group on voles and their prevalence was positively correlated with mean abundance (r = 0.73; P < 0.05). As a single rodent species, E. miletus has a high potential to harbour abundant ectoparasites with high species diversity and high rates of infestation. The sex of the vole affects ectoparasite infestation.     

Effect of a cotton intercrop on spider mite populations in jujube trees

In the arid and semi-arid parts of Xinjiang (China), jujube (Zizyphus jujube Mill) orchards are regularly intercropped with cotton. In 2016–2017, a field study was conducted to compare the species composition and infestation pressure of spider mites in local jujube trees under a cotton-jujube intercropping system and a jujube monocrop system. Our results showed that Tetranychus truncatus Ehara accounted for 94.2–98.5% of all spider mites in either cropping systems. Overall abundance of tree-inhabiting spider mites, including T. truncatus, in intercropped orchards was generally higher than in monocrop ones. Combining our work with earlier findings, we demonstrate how intercropping of cotton and jujube trees generally increases population levels of spider mites on either crop.     

Preliminary Molecular Characterizations of Sarcoptes scaibiei (Acari: Sarcoptidae) from Farm Animals in Egypt

Little is known about the genetic diversity of Sarcoptes scabiei mites in farm animals in Egypt. In this study, we characterized S. scabiei in 25 skin scrapes from water buffalo, cattle, sheep, and rabbits at the nuclear marker ITS2 and mitochondrial markers COX1 and 16S rRNA. Sequences of the ITS2 showed no host segregation or geographical isolation, whereas those of the mitochondrial COX1 and 16S rRNA genes indicated the presence of both host-adapted and geographically segregated populations of S. scabiei. Host adaptation may limit inter-species transmission of. S. scabiei, thus restrict gene flow among S. scabiei from different hosts. This is the first report on the molecular characterization of sarcoptic mites in Egypt. Further genetic studies involving larger numbers of specimens, especially those from humans and companion animals, are needed to understand the molecular epidemiology of sarcoptic mange in Egypt.     

Parasitism of the rhizocephalan <Emphasis Type="Italic">Briarosaccus callosus</Emphasis> on the crab <Emphasis Type="Italic">Paralomis verrilli</Emphasis> (Crustacea: Lithodidae) in the waters of southeast Sakhalin

The crab Paralomis verrilli in the waters of southeast Sakhalin is infected by the parasitic rhizocephalan barnacle Briarosaccus callosus. The prevalence of parasitic infection was on the average 4.36% (6.29% for females, 3.28% for males) and varied between samples from none up to 14.9%. The degree of prevalence was not related to the average carapace width and the sex ratio of crabs in samples. Sterilization of female P. verrilli was caused by the B. callosus infestation or its consequences. No more than two parasite externae per crab were found. Crabs with two externae made up 5.6% of all infected specimens. The infestation of crab hosts with two B. callosus externae negatively influenced the growth of the externae. A positive relationship was found between the width of the crab carapace and the length of the parasitic externae. The survival rate of P. verrilli with either one or two parasite B. callosus externae did not differ substantially. The proportion of crabs with externae and those with “scars” (12.2% in our case) can be taken as the index of survival of the parasitized crabs.     

Biology and host specificity of the genusPsoroptes Gervais (Acarina: Psoroptidae), with reference to its occurrence in Australia

Psoroptes mites are obligate ectoparasites of mammals that cause psoroptic mange in a number of domestic and wildlife species world-wide. Correct identification of species in this genus is important for eradication and quarantine programs. However, conflicting reports about species identity and host specificity within this morphologically conservative genus have raised questions regarding the usefulness of the taxonomic characters previously used to distinguish species. Records that rely only on host species do not provide accurate identification ofPsoroptes mites at the species level. This paper reviews the information available on life history, geographic distribution, and host specificity for this genus, with special reference to problems associated with identifying species in Australia.     

The occurrence of mites in cereal-based foods destined for human consumption and possible consequences of infestation

Seven categories of cereal-based food products purchased at food retail outlets in UK were examined for the presence of mites by analysing 20g samples using a flotation method. Mites were found in 21% of 571 samples, which were examined soon after purchase, and in 38% of 421 samples, derived from the 571 samples which were examined after 6 weeks of storage in volunteers' homes. Most of the samples where mites were detected had fewer than five mites. However, a few samples contained more than 20 mites with a maximum of 428 mites detected in a single sample. Sixteen families, genera or species of mites were recovered. The most common species were Acarus siro, Tyrophagus putrescentiae, Lepidoglyphus destructor and Glycyphagus domesticus. The level of contamination was broadly similar for each of the seven categories of products examined. It is likely that infestation occurs at every stage of food processing and storage. This is the first study of its kind and there is no reason to believe that results from similar studies carried out in other temperate countries would be markedly different. Increase in the percentage of contaminated samples following storage in a domestic situation suggests that the domestic environment is an important factor in developing infestation. The implications of mite contamination on food quality and on human health are briefly discussed.     

Distribution and Seasonal Abundance of Trematode Parasites (Trematoda: Allocreadiidae: <Emphasis Type="Italic">Crepidostomum</Emphasis> spp.) in burrowing-mayfly nymphs (Ephemeroptera: Ephemeridae: <Emphasis Type="Italic">Hexagenia</Emphasis> spp.) from Connecting Rivers of the Laurentian Great Lakes

Burrowing-mayfly nymphs such as Hexagenia spp. have been used extensively in North America and Europe as a biomonitoring tool to indicate mesotrophic water quality, yet infestation by associated parasites has not been well documented. We performed laboratory analysis of archived samples of Hexagenia spp. nymphs collected in 1985 and 1986 to provide base-line data on the distribution (1985) and seasonal infestation (1986) of the trematode parasite Crepidostomum spp. in Hexagenia spp. nymphs in connecting rivers between Lakes Superior and Erie of the Laurentian Great Lakes. In May and June 1985, frequency of occurrence of metacercariae was widely distributed throughout the connecting rivers (63% of 203 stations with nymphs), except in areas where nymph densities were relatively low (i.e.,≤69 nymphs/m²). Distribution was probably underestimated in the present study because of low probability (mean = 31%, range = 0–57%) of detecting infestation in a small number of collected nymphs ( ≤10) at nymph densities ≤69/m². In 1986, seasonal infestation between April and October occurred in 3.3% (627) of 18696 nymphs. Overall prevalence, mean intensity, and mean abundance of parasites at one station in the St. Marys River indicate parasite transmission occurred between June and September. This period of transmission is dependent on the life-cycle of the parasite. In addition, the life-cycle of Hexagenia spp. determines which annual cohort of nymphs is infested and therefore, the duration of infestation. Although, no impacts of infestation on Hexagenia spp. nymphs were observed in the present study, infestation intensities were high enough (≥25 metacercariae per nymph) at one station in the St. Marys River to potentially cause tissue damage in a high proportion (53%) of infested nymphs.     

Mitochondrial Genome Sequence of the Scabies Mite Provides Insight into the Genetic Diversity of Individual Scabies Infections

The scabies mite, Sarcoptes scabiei, is an obligate parasite of the skin that infects humans and other animal species, causing scabies, a contagious disease characterized by extreme itching. Scabies infections are a major health problem, particularly in remote Indigenous communities in Australia, where co-infection of epidermal scabies lesions by Group A Streptococci or Staphylococcus aureus is thought to be responsible for the high rate of rheumatic heart disease and chronic kidney disease. We collected and separately sequenced mite DNA from several pools of thousands of whole mites from a porcine model of scabies (S. scabiei var. suis) and two human patients (S. scabiei var. hominis) living in different regions of northern Australia. Our sequencing samples the mite and its metagenome, including the mite gut flora and the wound micro-environment. Here, we describe the mitochondrial genome of the scabies mite. We developed a new de novo assembly pipeline based on a bait-and-reassemble strategy, which produced a 14 kilobase mitochondrial genome sequence assembly. We also annotated 35 genes and have compared these to other Acari mites. We identified single nucleotide polymorphisms (SNPs) and used these to infer the presence of six haplogroups in our samples, Remarkably, these fall into two closely-related clades with one clade including both human and pig varieties. This supports earlier findings that only limited genetic differences may separate some human and animal varieties, and raises the possibility of cross-host infections. Finally, we used these mitochondrial haplotypes to show that the genetic diversity of individual infections is typically small with 1–3 distinct haplotypes per infestation.     

Co-evolution between ectoparasites and their insect hosts: a simulation study of a damselfly–water mite interaction

1. A simulation model investigating the co‐evolution of water mites infesting their aquatic insect hosts during emergence is presented. The model is based on field and experimental studies of the ectoparasitic water mite Arrenurus cuspidator and the damselfly Coenagrion puella. 2. Three scenarios were studied: (1) Only the host was allowed to evolve timing of emergence, while the timing of the parasites' infestation opportunity was held constant. (2) Both host and parasite were allowed to evolve. (3) Only the parasite's timing was allowed to evolve, while the host was constrained completely. 3. In the first two scenarios, parasite abundances decreased in the course of evolution and reached values well below those found in the field, whereas in the third scenario, parasite abundances were maintained at a level close to that found in the field. In the second scenario (co‐evolution), the host seemed to be the leader in the evolutionary race. 4. It is concluded that water mite parasitism is capable of shaping emergence patterns in aquatic insects and, despite the same life‐cycle length for host and parasite, the parasite evolves fast enough to shape its hatching pattern to match the emergence pattern of its host.