Host-associated Differentiation Research Articles

Host-associated genetic differentiation in the goldenrod elliptical-gall moth, Gnorimoschema gallaesolidaginis (Lepidoptera: Gelechiidae).

Careful study of apparently generalist phytophagous insects often reveals that they instead represent complexes of genetically differentiated host races or cryptic species. The goldenrod elliptical-gall moth, Gnorimoschema gallaesolidaginis, attacks two goldenrods in the Solidago canadensis complex: S. altissima and S. gigantea (Asteraceae). We tested for host-associated genetic differentiation in G. gallaesolidaginis via analysis of variation at 12 allozyme loci among larvae collected at six sites in Iowa, Minnesota, and Nebraska. Gnorimoschema gallaesolidaginis from each host are highly polymorphic (3.6-4.7 alleles/locus and expected heterozygosity 0.28-0.38 within site-host combinations). Although there were no fixed differences between larvae from S. altissima and S. gigantea at any site, these represent well differentiated host forms, with 11 of 12 loci showing significantly different allele frequencies between host-associated collections at one or more sites. Host plant has a larger effect on genetic structure among populations than does location (Wright's FST = 0.16 between host forms vs. F(ST) = 0.061 and 0.026 among altissima and gigantea populations, respectively). The estimated F(ST) between host forms suggests that the historical effective rate of gene flow has been low (N(e)m approximately 1.3). Consistent with this historical estimate is the absence of detectable recombinant (hybrid and introgressant between host form) individuals in contemporary populations (none of 431 genotyped individuals). Upper 95% confidence limits for the frequency of recombinant individuals range from 5% to 9%. Host association is tight, but imperfect, with only one likely example of a host mismatch (a larva galling the wrong host species). Our inferences about hybridization and host association are based on new maximum-likelihood methods for estimating frequencies of genealogical classes (in this case, two parental classes, F1 and F2 hybrids, and backcrosses) in a population and for assigning individuals to genealogical classes. We describe these new methods in the context of their application to genetic structure in G. gallaesolidaginis. Population phenograms are consistent with the origin of the host forms (at least in the midwestern United States) via a single host shift: altissima and gigantea moth populations form distinct lineages with 100% bootstrap support. Genetic structure in Gnorimoschema is of particular interest because another gallmaking insect attacking the same pair of hosts, the tephritid fly Eurosta solidaginis, includes a pair of host races with partial reproductive isolation. Gnorimoschema gallaesolidaginis and E. solidaginis therefore represent the first reported case of parallel host-associated differentiation, that is, differentiation by evolutionarily independent insect lineages across the same pair of host plants.

HOST-ASSOCIATED GENETIC DIFFERENTIATION IN THE GOLDENROD ELLIPTICAL-GALL MOTH, GNORIMOSCHEMA GALLAESOLIDAGINIS (LEPIDOPTERA: GELECHIIDAE)

Careful study of apparently generalist phytophagous insects often reveals that they instead represent complexes of genetically differentiated host races or cryptic species. The goldenrod elliptical-gall moth, Gnorimoschema gallaesolidaginis, attacks two goldenrods in the Solidago canadensis complex: S. altissima and S. gigantea (Asteraceae). We tested for host-associated genetic differentiation in G. gallaesolidaginis via analysis of variation at 12 allozyme loci among larvae collected at six sites in Iowa, Minnesota, and Nebraska. Gnorimoschema gallaesolidaginis from each host are highly polymorphic (3.6–4.7 alleles/locus and expected heterozygosity 0.28–0.38 within site-host combinations). Although there were no fixed differences between larvae from S. altissima and S. gigantea at any site, these represent well differentiated host forms, with 11 of 12 loci showing significantly different allele frequencies between host-associated collections at one or more sites. Host plant has a larger effect on genetic structure among populations than does location (Wright's FST = 0.16 between host forms vs. FST = 0.061 and 0.026 among altissima and gigantea populations, respectively). The estimated FST between host forms suggests that the historical effective rate of gene flow has been low (Nem ≈ 1.3). Consistent with this historical estimate is the absence of detectable recombinant (hybrid and introgressant between host form) individuals in contemporary populations (none of 431 genotyped individuals). Upper 95% confidence limits for the frequency of recombinant individuals range from 5% to 9%. Host association is tight, but imperfect, with only one likely example of a host mismatch (a larva galling the wrong host species). Our inferences about hybridization and host association are based on new maximum-likelihood methods for estimating frequencies of genealogical classes (in this case, two parental classes, F1 and F2 hybrids, and backcrosses) in a population and for assigning individuals to genealogical classes. We describe these new methods in the context of their application to genetic structure in G. gallaesolidaginis. Population phenograms are consistent with the origin of the host forms (at least in the midwestern United States) via a single host shift: altissima and gigantea moth populations form distinct lineages with 100% bootstrap support. Genetic structure in Gnorimoschema is of particular interest because another gallmaking insect attacking the same pair of hosts, the tephritid fly Eurosta solidaginis, includes a pair of host races with partial reproductive isolation. Gnorimoschema gallaesolidaginis and E. solidaginis therefore represent the first reported case of parallel host-associated differentiation, that is, differentiation by evolutionarily independent insect lineages across the same pair of host plants.

Genetic differentiation in Tetranychus urticae (Acari: Tetranychidae): polymorphism, host races or sibling species?

Based on allozyme electrophoresis at the Pgm locus and nuclear ribosomal DNA (ITS2) sequences, we studied the genetic variation of the two-spotted spider mite Tetranychus urticae Koch collected on rose bay, Nerium oleander L. (Apocynaceae), from several localities around the Mediterranean basin. In addition, we compared these results with those of Navajas et al. (1998) and Tsagkarakou (1997) who collected from several other host plants from the Mediterranean. In the western part of this area (Spain, France, Tunisia), we found the individuals collected from rose bay to be clearly genetically differentiated from other samples. No evidence of such host-associated differentiation was detected in the eastern Mediterranean (Italy and Greece). The genetic differentiation of mites collected on rose bay was investigated further by studying the reproductive incompatibilities between populations in Greece and in France and a laboratory strain reared on bean, Phaseolus vulgaris, in France. Reciprocal crosses performed between these strains revealed variable levels of incompatibility, spanning from partial to complete reproductive isolation. In all cases incompatibility was asymmetric. We designed a test based on double-mating to establish the fertilization status of females in fully incompatible crosses. These crosses showed that the females had been inseminated, which suggests that the barrier to reproduction is not of a prezygotic behavioral nature. The data raises the question of the relative role of ecological factors (host plant) and geographical distance, in the ongoing differentiation process potentially leading to speciation.

RESOURCE-ASSOCIATED POPULATION SUBDIVISION IN A SYMBIOTIC CORAL-REEF SHRIMP.

The importance of sympatric speciation remains controversial. An empirical observation frequently offered in its support is the occurrence of sister taxa living in sympatry but using different resources. To examine the possibility of sympatric differentiation in producing such cases, I measured genetic, behavioral, and demographic differentiation between populations of the tropical sponge-dwelling shrimp Synalpheus brooksi occupying two alternate host species on three reefs in Caribbean Panama. This species belongs to an apparently monophyletic group of ≥ 30 species of mostly obligate, host-specific sponge-dwellers, many of which occur in sympatry. Demographic data demonstrated the potential for disruptive selection imposed by the two host species: shrimp demes from the sponge Agelas clathrodes were consistently denser, poorer in mature females, more heavily parasitized by branchial bopyrid isopods, and less parasitized by thoracic isopods, than conspecific shrimp from the sponge Spheciospongia vesparium. Laboratory assays demonstrated divergence in host preference: shrimp on all three reefs tended to choose their native sponge species more often than did conspecific shrimp from the other host. Because S. brooksi mates within the host, this habitat selection should foster assortative mating by host species. A hierarchical survey of protein-electrophoretic variation also supported host-mediated divergence, revealing the following: (1) shrimp from the two hosts are conspecific, as evidenced by absence of fixed allelic differences at any of nine allozyme loci scored; (2) strong genetic subdivision among populations of this philopatric shrimp on reefs separated by 1-3 km; and (3) significant host-associated genetic differentiation within two of the three reefs. Finally, intersexual aggression (a proxy for mating incompatibility) between shrimp from different host species was significantly elevated on the one reef where host-associated genetic differences were strongest, demonstrating concordance between genetic and behavioral estimates of divergence. Adjacent reefs appear to be semi-independent sites of host-associated differentiation, as evidenced by differences in the degree of host-associated behavioral and genetic differentiation, and in the specific loci involved, on different reefs. In philopatric organisms with highly subdivided populations, such as S. brooksi, resource-associated differentiation can occur independently in different populations, thus providing multiple "experiments" in differentiation and resulting in a mosaic pattern of polymorphism as reflected by neutral genetic markers. Several freshwater fishes, an amphipod, and a snail similarly show independent but remarkably convergent patterns of resource-associated divergence in different conspecific populations, often in the absence of obvious spatial barriers. In each case, substantial differentiation has occurred in the face of continuing gene flow.

Geographic and Host-Associated Differentiation in the Fruittree Leafroller (Lepidoptera: Tortricidae)

The fruittree leafroller, Archips argyrospila (Walker), has recently reached out-break densities on a newly recorded host plant, baldcypress, Taxodium distichum (L.) Richard. We conducted a genetic analysis of the cypress population and one feeding on oaks in Louisiana. To obtain a geographic perspective on differentiation within the fruittree leafroller, we compared the Louisiana cypress population with a California oak population genetically and conducted cross-rearing studies on oak and cypress. A citrus population from California was also included for a host-associated perspective. Genetic data suggest the existence of three distinct taxa. One taxon feeds on oaks and citrus in California, one feeds on oaks in Louisiana, and a third feeds on baldcypress in Louisiana. Developmental cross-rearing studies indicate the cypress population is virtually unable to survive on Louisiana oaks, further supporting the existence of host-associated differentiation. Pheromone attraction studies conducted in Louisiana forests with California citrus and Louisiana cypress virgin females indicate California citrus females are not attractive to male fruittree leafrollers in cypress swamps. Differences in pheromonal response suggest evolutionary divergence in mating behavior on a geographic scale. Further studies are required to determine the degree of reproductive isolation between host associated and geographic populations, but genetic, developmental, and pheromonal studies suggest sibling species or host races, or both, exist.

The effects of climate, host plant phenology and host fidelity on the genetics of apple and hawthorn infesting races of Rhagoletis pomonella

AbstractTrue fruit flies belonging to the Rhagoletis pomonella (Walsh) sibling species complex have been proposed to speciate sympatrically by shifting and adapting to new host plants. Here, we report the results from a series of ecological and genetic experiments conducted at a study site near Grant, Michigan, U.S.A., aimed at clarifying the relationship between host specialization and reproductive isolation for these flies. Our findings indicate that apple (Malus pumila) and hawthorn (Crataegus mollis) infesting populations of R. pomonella are partially allochronically isolated. Differences in the timing of adult eclosion account for part of the allochronic divergence, as apple adults emerge approximately ten days earlier than hawthorn flies in the field. Genetic analyses across different life history stages of the fly show that adults do not randomly move between apple and hawthorn trees, but trend to attack the same species of plant that they infested as larvae. Estimates of interhost migration from the allozyme data suggest that from 2.8 to 10% of the apple population is of hawthorn origin and that over 20% of the hawthorn population is of apple origin. The length and quality of the growing season appear to affect the genetic composition of the host races, as allele frequencies in the hawthorn population are correlated with ambient temperature and rainfall during the spring of the preceding year. Finally, allele frequencies for six allozyme loci displaying host associated differentiation also show significant linear regressions with the timing of adult eclosion within both races. These regressions establish a link between allozyme loci displaying inter‐host differentiation and a developmental trait (adult eclosion) responsible for partially isolating the races. The slopes of the regressions are paradoxical, however, as they suggest that apple adults should eclose later, not earlier, than hawthorn flies. We conclude by discussing potential resolutions to the eclosion time paradox.

Nutritional performance of first and penultimate‐final Instar bagworm larvae on two unrelated hosts as influenced by host of origin

AbstractNutritional performance of early and late instar bagworm (Thyridopteryx ephemeraeformis Haworth) larvae was investigated as a function of host of origin and host type, using two taxonomically distant wild hosts, Salix nigra (Marshall) and Juniperus virginiana (L.). First instar growth and survival were higher on S. nigra; overall performance was similar on the two hosts in penultimate‐final instars. There was no evidence for host associated differentiation with respect to nutritional performance. There were, however, significant genotype & environment interactions for several characters studied, suggesting that the potential for host associated differentiation exists.

On the Origin of the Field Plants of the Northeastern United States

Old fields have dominated studies of terrestrial plant succession in the eastern United States. In spite of the tremendous amount of work on old field succession, fundamental questions remain. One such question is the subject of the present paper: In the primeval landscape, where were the plants that are abundant today in old fields? The question holds interest not only for interpretation of the northeastern landscape but also for understanding of the habitats in which field plants evolved. The plants of fallow agricultural fields are widely assumed to be early successional species, i.e., plants adapted to move from one temporary opening to another, but did they really evolve as successional plants? Many of the plants that grow in abandoned fields today are not native to North America and were thus not part of the landscape prior to European settlement. Bard (1952) reported that between one fifth and one third of the plant species in old fields in New Jersey were alien to North America. It is of the remaining two thirds (roughly 100 species) that we can ask, Where was a sun-loving field flora growing in a landscape that was heavily forested? According to Braun (1950, p. 7), "When the first settlers colonized the Atlantic seaboard ... they found an endless expanse of forest broken only by swamps and bogs, by cliffs or river bluffs too steep for forest, by windfalls and burns, by small grassy openings such as the serpentine barrens of the northern Piedmont, and by the prairie patches and 'barrens' of the interior.' Four hypotheses can be suggested to account for the location of native field plants in the presettlement landscape. (1) They evolved on agricultural land abandoned by Indians far back in time. (2) Native field plants migrated eastward from prairies and savannas at the western margin of the deciduous forest with the onset of extensive clearing for settlement and agriculture and with the development of continuous east-west corridors of open field habitat along railroads and canals. (3) Native field plants were true pioneer or early successional species that colonized temporary forest openings caused by wind, fire, and other kinds of