An investigation of the host range of the carrot fly

The historical records of the earliest reports of umbelliferous and other hosts of carrot fly ( Psila rosae ) are presented and their validity discussed. In addition, the results are reported of tests made between 1972 and 1980 on 27 species of Umbelliferae and three sub-species of Daucus carota to determine whether they would support the carrot fly. Techniques were developed for identifying plant species as hosts by determining the number of adult flies that emerged from root and soil samples obtained from plants exposed to carrot fly in the field. Experiments in 1979 and 1980 indicating the importance of co-ordinating the life cycles of plant and insect. The results showed that certain annual species of Umbelliferae were unsuitable as hosts in the autumn when they has flowered and senesced. Valid records could only be obtained for these species by exposing the plants to first generation carrot fly attack. Of the 27 species tested, Anthriscus sylvestris, Ferula communis, and Smyrnium olusatrum failed to support carrot fly and the following 13 species were new host records: Aethusa cynapium, Anthriscus caucalis, Apium inundatum, Apium nodiflorum, Bupleurum tenuissimum, Chaerophyllum temulentum, Cicuta virosa, Daucus capillifolius, Daucus glochidiatus, Heracleum mantegazzianum, Ligusticum scoticum, Oenanthe crocata and Sison amomum . From two species, Chaerophyllum temulentum and Torilis japonica , more than two carrot flies/root were recovered. These species are numerous and widespread in carrot-growing districts. The significance of these and other wild host plants in relation to the ecology and control of carrot fly is discussed.     

Host-plant susceptibility to the carrot fly, Psila rosae. 1. Acceptability of various host species to ovipositing females

A preference hierarchy of the oligophagous carrot fly for 30 umbelliferous host-plant species or varieties and six non-hosts was assessed. Foliage of the test plants was presented to a laboratory population of the fly in oviposition choice experiments together with leaves of a standard plant, the susceptible carrot cultivar “Danvers”. Only two species (Anthriscus cerefolium, Carum carvi) were more acceptable than the standard plant, while about half the species received significantly fewer eggs. Some umbellifers (Anethum graveolens, Pimpinella major) had a low acceptability similar to that of non-hosts (non-umbelliferous plants). The results obtained with dual and multiple choice assays were in agreement. A multiple choice assay with potted plants yielded a similar ranking of the species as assays using detached leaves, indicating that foliage is representative for whole plants. Variability in the exploratory runs performed by the females on the leaves prior to egg-laying is described and quantified for hosts and non-hosts. Differences in post-alighting pre-ovipositional behaviour of the female flies on the leaves accounted for a major part of the variation recorded in egg-laying.     

Host-plant susceptibility to the carrot fly, Psila rosae. 2. Suitability of various host species for larval development

Several apiaceous and two asteraceous species were tested for their suitability to support larval development of the carrot fly. Plants grown in pots or transplanted from seed beds into pots, were inoculated with a specific number of eggs. Pupae and non-pupated larvae were collected 6–7 wk after inoculation. Both the number and weights of pupae produced varied widely among the species. Cultivated carrots Daucus carota sativus often gave rise to only moderate numbers of pupae, but these invariably attained the highest weights. Pimpinella major was the only apiaceous plant tested that did not yield any carrot flies. The two asteraceous plants Cichorium intybus and Tanacetum vulgare failed to support larval development. Total carrot fly biomass produced per plant was influenced by both the host species and the root weight. Emergence rates of adult flies were positively correlated with pupal weights. Small individuals tended to have a longer total developmental time from egg to adult fly.     

Monitoring populations of the carrot fly, Psila rosae, with sticky and water traps

The relative, effectiveness of Rebell®, small cylinder, large cylinder, windmill and water traps, the five types of trap used currently for monitoring populations of the carrot fly, Psila rosae, was assessed in nine field experiments, three in south west Lancashire, four in the Fens (Suffolk, Norfolk, Cambridgeshire), one in East Suffolk and one at Wellesbourne, Warwickshire. Regression analysis of the numbers of flies caught on each type of trap against the numbers caught on the Rebell® trap indicated that each trap samples a constant proportion of the fly population relative to the other traps. Therefore, provided the fly population was sufficiently large for insects to be caught on the least effective traps, any of the five traps would monitor adequately fluctuations in carrot fly populations. However, the Rebell® trap caught 4–17 times as many flies/trap and 5–7 times as many flieshnit area of trap as any of the other traps tested.
Operators considered the Rebell® trap to be the easiest to use. However, it was more expensive than any of the other traps tested.     

Exploitation of the resistance to carrot fly in the wild carrot species Daucus capillifolius

Under field conditions a wild Daucus species from Libya, D. capillifolius, supported less than one tenth as many carrot flies (Psila rosae) as the susceptible carrot cultivar Danvers Half Long 126. Breeding lines developed from crosses between D. capillifolius and three different carrot types were grown in a series of field experiments at Wellesbourne between 1980 and 1989. Each year selections were made for agronomic quality and/or for increased resistance to carrot fly. The programme produced lines which for size, shape and colour represented most of the commercially-important carrot types. Some of these lines were also significantly more resistant to carrot fly than selections from the partially-resistant cv. Sytan. However, the best lines were not as resistant as the wild parent. The highest quality resistant lines were sold to seed companies for variety production.     

The occurrence of Entomophthorales on the carrot fly [Psila rosae F.] in the field during two successive seasons

Three species of Entomophthorales were found on adult carrot flies (Psila rosae F.) during two successive seasons:Entomophthora muscae (C.) Fres.,Condiobolus apiculatus (Thax.) Remaud. & Keller andErynia sp.E. muscae was by far the most common species and caused epizootics in one carrot fly population up to 3 times per year. Flies cuaght in the hedge showed a higher infection level than flies caught in the field close to the hedge. Flies caught 200 m away from the hedge showed the lowest infection level. Apparently the hedge was the site of infection, since carrot flies, killed byE. muscae, were found there attached to the underside of the leaves. Flies caught on yellow sticky traps developed only to a limited extend symptoms and gave little information about the fungus infection levels.     

Studies on the aphid, Cavariella aegopodii Scop: On secondary hosts other than carrot

Of the eleven cultivated species of Umbelliferae heavily attacked in the field by Cavariella aegopodii Scop., chervil (Anthriscus cerefolium) attracted so many alates that it was killed before a population of alienicolae developed. Carrot was next in order of alate preference and coriander (Coriandrum sativum) and caraway (Carum carvi) were last. Despite its lack of attraction, caraway proved more lastingly suitable as a host and eventually supported the largest population of alienicolae. Of the wild umbellifers tested, wild carrot, Daucus carota ssp. carota, was the species most preferred by the incoming alates and hemlock (Conium maculatum) the least. On hemlock, also, there was little colonization. Although a sequence of wild umbellifers supported C. aegopodii through most of the year, few were suitable winter hosts; the apterae hibernated largely on carrot crops. In spring, cow parsley (Anthriscus sylvestris) was one of the hosts first colonized by immigrant C. aegopodii and carried winged alienicolae before the end of June. The main development of winged alienicolae on Anthriscus and other wild Umbelliferae, however, was in July. The possible role of the wild umbelliferous hosts of C. aegopodii in the spread of motley dwarf virus to carrots is discussed.     

Olfactory responses of the omnivorous generalist predator Dicyphus hesperus to plant and prey odours

This paper describes a method for studying the neighbourhood (dispersal) movement of pest insects that overcomes the major problems associated with release/recapture experiments. The method has been developed using the carrot fly, Psila rosae (Fab.) (Diptera: Psilidae), as the experimental insect. It involves building up a large population of insects in an area free of other host plants, and then, as the insects disperse in the spring, monitoring the fall‐off in numbers of insects with increasing distance. Estimates of the fall‐off in numbers of the carrot fly were obtained by using sticky traps to monitor changes in fly numbers in 11 small plots of carrots drilled 130–1300 m away from the site at which the flies emerged (emergence site) in the spring. A strong linear relationship (r² = 0.96) was obtained between the log₁₀ numbers of flies caught in each plot and the log₁₀ distance the plot was from the emergence site. Few flies were caught more than 1 km from the emergence site. A linear relationship (r² = 0.86) was also obtained between the date when 50% of the flies were caught in each plot and the distance the plot was from the emergence site. The date of 50% capture was delayed by 1 day for each 100 m the given plot was away from the emergence site. This suggests that when carrot flies move to find new crops, the population moves about 100 m day^?1. About one million flies would have to be released in conventional release/recapture experiments to obtain data as robust as those collected using the current method. The major breakthrough of this method is that dispersal can be estimated for an insect species that, although a major pest of field crops of carrots and parsnips, is exceptionally difficult to rear, even in relatively small numbers, under controlled‐environment conditions.     

Inhibition of the DNA amplification of trypanosomes present in tsetse flies midguts: implications for the identification of trypanosome species in wild tsetse flies

The present study was carried out in order to investigate if there was really a failure of PCR in identifying parasitologically positive tsetse flies in the field. Tsetse flies (Glossina palpalis gambiensis and Glossina morsitans morsitans) were therefore experimentally infected with two different species of Trypanosoma (Trypanosoma brucei gambiense or Trypanosoma congolense). A total of 152 tsetse flies were dissected, and organs of each fly (midgut, proboscis or salivary glands) were examined. The positive organs were then analysed using PCR. Results showed that, regardless of the trypanosome species, PCR failed to amplify 40% of the parasitologically positive midguts. This failure, which does not occur with diluted samples, is likely to be caused by an inhibition of the amplification reaction. This finding has important implications for the detection and the identification of trypanosome species in wild tsetse flies.     

Community of bat flies (Streblidae and Nycteribiidae) on bats in the Cerrado of Central-West Brazil: hosts,aggregation, prevalence,infestation intensity,and infracommunities

Streblidae and Nycteribiidae are families of bloodsucking flies that parasitize bats exclusively. We studied the community of these flies in a Cerrado area in the Central-West Brazil. We captured 708 bats over 17 nights from October 2012 to March 2013. Forty-five per cent of the hosts were parasitized by 836 specimens of bat flies of 22 species. The most abundant flies were Trichobius joblingi on Carollia perspicillata, followed by Megistopoda aranea on Artibeus planirostris, and Strebla guajiro on C. perspicillata. All bat flies showed a high level of specificity for their hosts. Trichobius joblingi was the bat fly with the highest prevalence (80%) and mean intensity of infestation (3.5) on hosts with a representative sample size (n > 20). This result is likely related to the type of roosting (cavity) used by C. perspicillata, primary host of this fly species. Anoura caudifer hosted the largest infracommunities (n = 7). However, most bats were parasitized by a single fly species, suggesting a pattern in infestations. The aggregation index was high, indicating an unequal occurrence in parasite infestations. The majority of hosts were infested by few or no flies and few hosts were highly infested, showing a negative binomial distribution.     

High hemocyte load is associated with increased resistance against parasitoids in Drosophila suzukii, a relative of D. melanogaster

Among the most common parasites of Drosophila in nature are parasitoid wasps, which lay their eggs in fly larvae and pupae. D. melanogaster larvae can mount a cellular immune response against wasp eggs, but female wasps inject venom along with their eggs to block this immune response. Genetic variation in flies for immune resistance against wasps and genetic variation in wasps for virulence against flies largely determines the outcome of any fly-wasp interaction. Interestingly, up to 90% of the variation in fly resistance against wasp parasitism has been linked to a very simple mechanism: flies with increased constitutive blood cell (hemocyte) production are more resistant. However, this relationship has not been tested for Drosophila hosts outside of the melanogaster subgroup, nor has it been tested across a diversity of parasitoid wasp species and strains. We compared hemocyte levels in two fly species from different subgroups, D. melanogaster and D. suzukii, and found that D. suzukii constitutively produces up to five times more hemocytes than D. melanogaster. Using a panel of 24 parasitoid wasp strains representing fifteen species, four families, and multiple virulence strategies, we found that D. suzukii was significantly more resistant to wasp parasitism than D. melanogaster. Thus, our data suggest that the relationship between hemocyte production and wasp resistance is general. However, at least one sympatric wasp species was a highly successful infector of D. suzukii, suggesting specialists can overcome the general resistance afforded to hosts by excessive hemocyte production. Given that D. suzukii is an emerging agricultural pest, identification of the few parasitoid wasps that successfully infect D. suzukii may have value for biocontrol.     

Nested distributions of bat flies (Diptera: Streblidae) on Neotropical bats: artifact and specificity in host-parasite studies

We examined the structure of ectoparasitic bat fly infestations on 31 well‐sampled bat species, representing 4 Neotropical families. Sample sizes varied from 22 to 1057 bats per species, and bat species were infested by 4 to 27 bat fly species. Individual bats supported smaller infracommunities (the set of parasites co‐occurring on an individual host), ranging from 1 to 5 fly species in size, and no bat species had more than 6 bat fly species characteristically associated with it (its primary fly species). Nestedness analyses used system temperature (BINMATNEST algorithm) because it is particularly well‐suited for analysis of interaction networks, where parasite records may be nested among hosts and host individuals simultaneously nested among parasites. Most species exhibited very low system temperatures (mean 3.14°; range 0.14–12.28°). Simulations showed that nested structure for all 31 species was significantly stronger than simulated values under 2 of the 3 null hypotheses, and about half the species were also nested under the more stringent conditions of the third null hypothesis. Yet this structure disappears when analyses are restricted to “primary” associations of fly species (flies on their customary host species), which exclude records thought to be atypical, transient, or potential contaminants. Despite comprising a small fraction of total parasite records, such anomalies represent a considerable part of the statistical state‐space, offering the illusion of significant ecological structure. Only well understood and well documented systems can make distinctions between primary and other occurrence records. Generally, nestedness appears best developed in host‐parasite systems where infestations are long‐term and accumulate over time. Dynamic, short‐term infestations by highly mobile parasites like bat flies may appear to be nested, but such structure is better understood in terms of host specificity and accidental occurrences than in terms of prevalence, persistence, or hierarchical niche relations of the flies.     

Electroantennogram responses of the carrot fly, Psila rosae, to volatile plant components

ABSTRACT. Electroantennogram (EAG) responses of male and female carrot flies, Psila rosae F. (Diptera: Psilidae), were recorded to thirty-six volatile plant constituents. The most distinct EAG responses were obtained to: (1) the general green leaf volatiles 1-hexanol, trans-2-hexen-1-ol and cis-3-hexen-1-ol, their isomers cis-2-hexen-1-ol and trans-3-hexen-1-ol, the alcohol 1-heptanol, the ester cis-3-hexenyl acetate and the leaf aldehydes hexanal and trans-2-hexenal, and (2) from four compounds associated with the umbelliferous host plants of this insect, namely trans-methyl-iso-eugenol, β-caryophyllene, linalool and trans-2-nonenal. Higher responses were elicited by the leaf aldehydes than by the corresponding alcohols. Although the absolute amplitude of the female response was over twice that of the male, there were no differences between the relative responses to the compounds tested in both sexes, with the exception of a much higher response to the leaf aldehydes in the male. The shape of the EAG evoked by the various compounds was consistently different, with the slowest recovery being recorded for trans-methyl-iso-eugenol. While the antennal olfactory receptors of the carrot fly are sensitive to the closely related general green leaf volatiles, they are most specifically tuned to the aldehyde component of this green odour complex. In addition, the ability of this insect to discriminate between different plants may be augmented by the perception of a group of more host specific volatiles. The conformity of the responses of males and females to the compounds tested may indicate that host plant volatiles plays an additional role as an aggregation cue for both sexes.     

Identification of the intermediate hosts of Habronema microstoma and Habronema muscae under field conditions

Abstract.  A polymerase chain reaction (PCR)-based assay was used for the specific detection of Habronema microstoma and Habronema muscae (Nematoda, Spirurida) in order to identify the intermediate hosts of both nematode species under field conditions. A total of 1087 netted and 165 laboratory-bred flies were tested. Flies were identified as Musca domestica Linnaeus 1758, Musca autumnalis De Geer 1776, Haematobia irritans (Linnaeus 1758), Haematobia titillans (De Geer 1907) and Stomoxys calcitrans (Linnaeus 1758) (Muscidae). Genomic DNA was extracted from pools of fly heads, thoraces and abdomens, and 703 samples were subjected to a duplex two-step semi-nested PCR assay to specifically detect diagnostic regions within the ribosomal ITS2 sequence of both H. microstoma and H. muscae . Stomoxys calcitrans specimens were positive for H. microstoma DNA and M. domestica specimens were positive for H. muscae DNA. In particular, PCR-positive samples derived from both farm-netted and laboratory-bred flies. The present study represents the first evidence of the vectorial competence of different fly species as intermediate hosts of Habronema stomachworms under field conditions. We discuss the roles of S. calcitrans and M. domestica in transmitting H. microstoma and H. muscae .     

Breeding sites of Phlebotomus sergenti, the sand fly vector of cutaneous leishmaniasis in the Judean Desert

Phlebotomine sand flies transmit Leishmania, phlebo-viruses and Bartonella to humans. A prominent gap in our knowledge of sand fly biology remains the ecology of their immature stages. Sand flies, unlike mosquitoes do not breed in water and only small numbers of larvae have been recovered from diverse habitats that provide stable temperatures, high humidity and decaying organic matter. We describe studies designed to identify and characterize sand fly breeding habitats in a Judean Desert focus of cutaneous leishmaniasis. To detect breeding habitats we constructed emergence traps comprising sand fly-proof netting covering defined areas or cave openings. Large size horizontal sticky traps within the confined spaces were used to trap the sand flies. Newly eclosed male sand flies were identified based on their un-rotated genitalia. Cumulative results show that Phlebotomus sergenti the vector of Leishmania tropica rests and breeds inside caves that are also home to rock hyraxes (the reservoir hosts of L. tropica) and several rodent species. Emerging sand flies were also trapped outside covered caves, probably arriving from other caves or from smaller, concealed cracks in the rocky ledges close by. Man-made support walls constructed with large boulders were also identified as breeding habitats for Ph. sergenti albeit less important than caves. Soil samples obtained from caves and burrows were rich in organic matter and salt content. In this study we developed and put into practice a generalized experimental scheme for identifying sand fly breeding habitats and for assessing the quantities of flies that emerge from them. An improved understanding of sand fly larval ecology should facilitate the implementation of effective control strategies of sand fly vectors of Leishmania.     

Comparison of the sensitivity of four Delia species to host and non-host plant compounds

Abstract. The behavioural facilitation hypothesis, tested in the present study, suggests that evolution of host-plant shifts by phytophagous insects is based on the preadaptation of insects to the chemistry of potentially novel plant hosts. Thus, closely-related insects should have similar sensitivities to compounds that are shared by different host plants. The chemoreception is investigated for four phytophagous flies, Delia radicum, Delia floralis, Delia antiqua and Delia platura (Diptera, Calyptratae: Anthomyiidae), belonging to the same genus but developing mainly on different plant families, with particular secondary plant compound profiles. In addition, the carrot fly, Psila rosae, an acalyptrate Diptera, is included as an unrelated species that is associated with completely different host plants. For the comparison, the known oviposition stimulants of the cabbage root fly (glucobrassicin, sinalbin, sinigrin and a thia-triaza-fluorene compound; CIF-1) present on the cabbage leaf surface were chosen. Responses from prothoracic tarsal sensilla are recorded to contact stimulation in a dose-dependent manner. Among the different flies tested, only D. radicum responds to all the compounds. By contrast, D. floralis is only sensitive to CIF-1, and not specifically on the C₅ sensillum, a finding that is in conflict with previously published results. This discrepancy is possibly an indication of the variability among flies originating from different cultures or habitats. With the exception of sinigrin at high concentration, the various compounds tested do not stimulate D. antiqua or D. platura. However, the carrot fly appears to be completely insensitive to sinigrin even at the highest tested concentration of 10⁻¹ M. The responses of the contact-chemoreceptor neurones to the selected compounds therefore provide little evidence of common sensitivities that would explain host shift in Delia species and specialization at the physiological level. The wide divergence within closely-related species and rearing cultures appears to indicate that the sensitivity and distribution of sensory receptor neurones is very variable on an evolutionary scale.     

A test of host specialization by insect vectors as a mechanism for reproductive isolation among entomophilous fungal species

Epichloë species are self incompatible (heterothallic) fungi that must be fertilized by spermatia from individuals of opposite mating type for successful sexual reproduction to occur. Female flies of the genus Botanophila act as vectors of the fungi by ingesting and defecating spermatia (gametes) onto fungal stromata (fruiting bodies) after oviposition. Larvae feed and develop on the stromata and thus maintain a symbiotic relationship with Epichloë fungi. We hypothesized that sole dependence on fertilized stromata as a food source would promote specialization by flies to single compatible host species and that this specialization would promote reproductive isolation among Epichloë species. Analysis of progeny of ascospores from experimental field plots in Zurich, Switzerland, indicated prevalence of specific matings between stromata of the same host, and thus was consistent with the hypothesis that flies are species-specific in their visitation behaviour. Genetic analyses of spermatia contained in the faeces of individual flies also gave some support for this hypothesis. We recovered spermatia of 4 different Epichloë species from fly faeces. Comparison of spermatia found in fly faeces to those available from stromata showed flies avoided Epichloë clarkii and may have preferred Epichloë typhina . Interestingly, these are the only two Epichloë species known to be interfertile with one another. Individual flies tended to carry spermatia predominantly from one fungal species. Thus, flies may adopt a type of "majoring" and "minoring" behaviour when visiting fungi. Yet, Botanophila flies are not monolectic and often visited all hosts that were available within screened cages. In addition to any reproductive isolation flies may provide to some fungal species, differences in competitiveness among spermatia of different species deposited on the same stroma may favor intraspecific matings.     

Microbial mediation of fruit fly–host plant interactions: is the host plant the “centre of activity”?

Insects utilize resources in their environment with the aid of mutualistic or symbiotic mediation by microorganisms. Some insect species such as ants and termites often have complex ecological and evolutionary associations with their symbionts, while the nature and functional significance of such associations in non-social insects is often unclear. In the Dacinae (Diptera: Tephritidae), specific Enterobacteriaceae ( Erwinia herbicola , Enterobacter cloacae , Klebsiella oxytoca ) are believed to mediate interactions between the adult fruit flies and the larval host plant. This bacterial mediation is hypothesized as being integral to the larval host plant being the "centre of activity" of the fly. Using a non-pest, monophagous fruit fly ( Bactrocera cacuminata [Hering]), we tested this hypothesis by manipulating the fruiting state of its larval host plant ( Solanum mauritianum Scopoli) and subsequently assessing insect behaviour and phylloplane microflora on those hosts. On host plants that had never fruited, few flies or bacterial colonies were recorded, consistent with hypothesis expectations. On fruiting host plants or plants that had had their fruit removed, bacterial colonies were present; again consistent with expectation. However, few flies were recorded on fruit-removed plants and all fly behaviours, other than resting or oviposition, were rare or absent on any hosts; inconsistent with expectation. The general pattern of results suggested that female flies coming to oviposit on fruiting hosts were spreading Enterobacteriaceae, but such spread was incidental and not part of some mutualistic interaction between fruit flies and bacteria.     

A hitchhiker’s guide to parasite transmission: The phoretic behaviour of feather lice

Transmission to new hosts is a fundamental challenge for parasites. Some species meet this challenge by hitchhiking on other, more mobile parasite species, a behaviour known as phoresis. For example, feather-feeding lice that parasitise birds disperse to new hosts by hitchhiking on parasitic louse flies, which fly between individual birds. Oddly, however, some species of feather lice do not engage in phoresis. For example, although Rock Pigeon (Columba livia) “wing” lice (Columbicola columbae) frequently move to new hosts phoretically on louse flies (Pseudolynchia canariensis), Rock Pigeon “body” lice (Campanulotes compar) do not. This difference in phoretic behaviour is puzzling because the two species of lice have very similar life cycles and are equally dependent on transmission to new hosts. We conducted a series of experiments designed to compare the orientation, locomotion and attachment capabilities of these two species of lice, in relation to louse flies. We show that wing lice use fly activity as a cue in orientation and locomotion, whereas body lice do not. We also show that wing lice are more capable of remaining attached to active flies that are walking, grooming or flying. The superior phoretic ability of wing lice may be related to morphological adaptations for life on wing feathers, compared to body feathers.     

Fitness effects of sex ratio response to host quality and size in the parasitoid wasp Spalangia cameroni

The parasitoid wasp Spalangia cameroni oviposited a greaterproportion of daughters in stable fly pupae than in house flypupae, even when I controlled for stable flies being smallerthan house flies. Sex ratio manipulation in response to hostquality has been modeled as being adaptive through an effectof host quality on the size and hence offspring production ofdaughters. 5. cameronis response to host species may insteadbe adaptive through an effect on larval survivorship, the developmenttime of daughters, and the size of sons. There was greater survivalof daughters than sons on stable flies. Controlling for hostsize, I found that development time of daughters was about 2%less on stable flies than on house flies. The decrease in developmenttime corresponds to a 2% increase in fitness as estimated byr, the intrinsic rate of increase, and is equivalent to abouta 9% increase in offspring production. Sons were about 2% largerfrom house flies than stable flies, which may increase offspringproduction by up to 3%. Host species had no consistent effecton size of daughters or development time of sons. In additionto the response to host species, mothers oviposited a greaterproportion of daughters in larger stable fly hosts. Whetherthis behavior is adaptive is unclear. Although offspring werelarger when they developed on larger stable flies, the rateof increase was less for daughters dian for sons. Effects ofstable fly size on offspring development time were negligible.