RANGE EXPANSION OF GREEN TREEFROGS (HYLA CINEREA) IN SOUTHERN ILLINOIS: NO EVIDENCE OF PARASITE RELEASE.

For several decades, green treefrogs (Hyla cinerea) have been undergoing rapid range expansion northward and eastward in Illinois, Indiana, and Kentucky. While range expansion of green treefrogs in these states may be linked to climate change, a recent study suggested this expansion could be facilitated by parasites, given that expanded range populations of green treefrogs from Kentucky and Indiana exhibited significant decreases in helminth species diversity compared to those examined from historical locations of Kentucky. Because rapid range expansion may lead to hosts escaping their parasites (= parasite release), a reprieve from parasitic infection could allocate additional resources to growth and reproduction and thus facilitate the expansion. The present study compares patterns of helminth diversity for green treefrogs from historical and 2 types (early and late expansion) of expanded range locations of southern Illinois to test whether these range-expansion populations are also experiencing a reduction in parasitism due to parasite release. The results of this study did not find significant differences in helminth diversity when helminth communities of green treefrogs from their historical and expanded ranges were compared. These results appear to downplay the putative role of parasite release in the northward range expansion of H. cinerea in Illinois. Studies are underway to determine whether local factors, including abiotic conditions and amphibian host diversity, play a more prominent role in influencing helminth diversity of green treefrogs.

HELMINTH DIVERSITY OF GREEN TREEFROGS (HYLA CINEREA) IN THEIR EXPANDED GEOGRAPHICAL RANGE.

There has been a growing interest in characterizing the parasite faunas of species populations as they expand their geographical ranges as a result of climate change. Expanded-range populations often exhibit lower parasite diversity than historical-range populations, and reduced parasitism may, in part, be attributable to expanded-range populations escaping their native range parasites. The present study compares the helminth faunas of green treefrogs (Hyla cinerea) from 4 historical and 4 expanded-range populations to determine whether these latter populations have undergone parasite escape. Results of this study found relatively high degrees of similarity in species composition among helminth assemblages within historical or within range-expansion locations, with marked differences in the composition of helminth faunas between historical and expanded-range populations of these frogs. Because green treefrogs from expanded-range locations exhibited significant decreases in helminth species diversity compared with those from historical sites, they appear to be escaping levels of parasitism typically experienced by these frogs in their native range. Most notably, there was a decrease in the abundance of helminths with direct life cycles and the absence of trematode assemblages with indirect life cycles among expanded-range populations of H. cinerea. The low prevalence of trematode assemblages among historical populations of green treefrogs could limit these parasites' ability to be introduced and propagated in expanded-range locations. However, the lack of trematode assemblages among populations of H. cinerea in its expanded range may also be due to the absence or limited availability of other aquatic hosts that are required to complete the life cycles of these parasites. The reduction in helminth diversity among expanded-range populations of green treefrogs lends some credence to the notion that individuals at the front of a range expansion often invest less energy in reproduction and in doing so allocate more energy to dispersal and other life-history traits, including resistance to parasites. There may, however, be other explanations for differences in parasite species diversity between historical and expanded-range populations of H. cinerea. Because many of the helminths reported from this study are host generalists of amphibians whose recruitment and transmission among intermediate and paratenic hosts are known to be constrained by water and/or soil moisture conditions, we cannot ignore the role of both local amphibian diversity and local abiotic factors in influencing helminth diversity between the 2 population types of green treefrogs. These latter factors would decrease the role of parasite escape or energy trade-offs in driving helminth diversity among populations of H. cinerea and instead would suggest that local conditions play a more prominent role in structuring their helminth communities.

Host Diversity Affects Parasite Diversity: A Case Study Involving Unionicola spp. Inhabiting Freshwater Mussels.

Water mites of the genus Unionicola are common parasites of freshwater mussels, living on the gills or mantle of their hosts and using these tissues as sites of oviposition. Although surveys of this mite fauna among North American unionid mussels indicate that these mites represent highly diverse assemblages, we know very little regarding the determinants of Unionicola species diversity among their molluscan hosts. The present study addresses the relationship between host diversity and mite diversity for Unionicola assemblages associated with unionid mussels of North America. The results of this study found a significantly positive relationship between host species richness and mite species richness, adding to a growing body of evidence that host diversity is an important determinant of parasite diversity. In recent years, molecular sequence data have discovered cryptic biodiversity among unionid mussels, yielding revisions in the nomenclature and systematic taxonomy of the group. DNA sequence variation has also revealed cryptic species complexes among Unionicola mites. Collectively, these findings suggest that the results of the present study may be underestimating species richness among mites and their host mussels. Unfortunately, human perturbations are known to have caused high recent rates of extinction in the mussel and mite faunas of North America and could play a major role in influencing patterns of species richness for this host-parasite system moving forward.

A Comparison of Helminth Faunas of Cope's Gray ( Hyla chrysoscelis ) and Green ( Hyla cinerea ) Treefrogs in Areas of Recent Niche Overlap.

This study compares the helminth faunas between Cope's gray treefrogs ( Hyla chrysoscelis ) and green treefrogs ( Hyla cinerea ), in areas where they have recently overlapped due to range expansion by H. cinerea , to determine whether or not 2 species of frogs with a high degree of similarity in many of their life history traits also exhibit similarities in the composition of their helminth assemblages. Results of this study did not find significant differences in helminth species diversity when sympatric and allopatric populations of the same species of frog were compared. There was, however, a significant difference in helminth diversity among sympatric populations of H. chrysoscelis and H. cinerea , and this difference was in large part attributable to the significantly higher abundance of the gastrointestinal nematode Cosmocercoides variabilis among H. chrysoscelis . Additional studies will be required to determine whether the observed patterns are due to differences in arrival time, perch locations within the chorus, or parasite-mediated competition.

Patterns of species richness among assemblages of Unionicola spp. (Acari: Unionicolidae) inhabiting freshwater mussels (Bivalvia: Unionoida) of North America.

Water mites of Unionicola species are common symbionts of freshwater mussels, living on the gills or mantle and foot of their hosts and using these tissues as sites of oviposition. Although surveys of the mite fauna among North American mussels suggest that these mites represent highly diverse assemblages, there are currently no quantitative data characterizing Unionicola species diversity among their molluscan hosts. The present study addresses patterns of species richness of Unionicola assemblages from freshwater mussels, including the relationship between richness and host specificity among these mites. Results from this study indicate that mite species richness increased significantly with an increase in the number of host individuals sampled. When corrected for sampling effort, there was a positive relationship between host size and mite species richness. Results from this study also reveal a significant relationship between mite species richness and the geographical distribution of host mussels. Overall, the patterns of species richness observed for this study are consistent with those examining the richness of parasitic helminth communities. Because the phylogenetic history of host taxa can have a significant influence on patterns of parasite species richness, studies that correct for the phylogenetic history among host mussels will be required to better understand the role that evolutionary processes have in determining Unionicola species richness. The present study did not indicate a significant relationship between species richness and host specificity and, in not doing so, suggests that the dispersal ability of mites may also play a role in influencing Unionicola species richness. The host recognition behavior and swimming abilities for a larger sample of mites will be required to substantiate this hypothesis.

Mitochondrial genome sequence of Unionicola parkeri (Acari: Trombidiformes: Unionicolidae): molecular synapomorphies between closely-related Unionicola gill mites.

The mitochondrial genome of Unionicola parkeri is a 14,734 bp circular DNA molecule. The sequence and annotation revealed a unique gene order, related to but distinct from the gene order in the closely related species U. foili. Mitochondrial tRNA sequences annotated in this genome predict non-canonical secondary structures for these molecules. The continuing patterns of unique gene orders and unusual tRNA structures in the Trombidiformes in general and Unionicola in particular support the use of phylogenetic approaches that use these types of molecular features as shared, derived character states. Further progress in using these molecular character states to reconstruct phylogeny will depend on careful annotation, especially of tRNA genes.

Mitochondrial genome sequence of Unionicola foili (Acari: Unionicolidae): a unique gene order with implications for phylogenetic inference.

The mitochondrial genome of Unionicola foili is circular, 14,738 bp in length, and contains several notable features. The sequence and annotation revealed a unique gene order, continuing a pattern of highly rearranged mitochondrial genomes in the Trombidiformes. U. foili mitochondrial tRNA sequences predict non-canonical secondary structures for these molecules, and our annotation suggests an in-frame fusion between the nad4L and nad5 genes in this genome. The unique gene order and unusual tRNA structures could serve as idiosyncratic characters and have the potential to be phylogenetically informative.

Host specificity among Unionicola spp. (Acari: Unionicolidae) parasitizing freshwater mussels.

Water mites of Unionicola spp. are common parasites of freshwater mussels as adults, living on the gills, or mantle and foot of their hosts and using these tissues as sites of oviposition. The present study addresses specialization among North American Unionicola mussel-mites using 2 measures of host specificity: (1) the number of host species used by a species of mite; and (2) a measure that considers the taxonomic distinctness of the hosts utilized by mites, weighted for their prevalence in the different hosts. Results of this study indicate the Unionicola spp. mussel-mites are highly host specific, with most species occurring in association with 1 or 2 species of hosts. If 2 or more host species are utilized, they are typically members of the same genus. These data are consistent with studies examining the dispersal abilities and host recognition behavior for members of the group. When the average values of host specificity for Unionicola subgenera were mapped on a phylogenetic tree for these taxa, a clade comprised of gill mites appeared to be more host specific than a clade consisting of mantle mites. There were, however, no apparent patterns of host specificity within each of the clades. Differences in specificity between the 2 lineages may reflect either a long evolutionary history that gill mites have had with host mussels or the intense competition among gill mites for oviposition sites within unionid mussels, leading to increased host specialization.

Random amplified polymorphic DNA analysis of kinship within host-associated populations of the symbiotic water mite Unionicola foili (Acari: Unionicolidae).

Kinship relations within populations of unionicolid water mites are not well known, owing to their complex life cycles and the fact that interactions between active and resting stages for some species are transitory. A number of species of unionicolid water mites are, however, obligate symbionts of freshwater mussels and spend most of their life cycle in association with these hosts. Among these species of mites, parents and offspring are more likely to co-occur and thus provide opportunities to address questions related to the structure of the mating system. The present study employs random amplified polymorphic DNA (RAPD) analysis to address kinship within populations of Unionicola foili living in symbiotic association with the host mussel Utterbackia imbecillis. DNA was amplified from adult mites and a representative number of eggs or larvae (n = 20-30) that were removed from mussels collected on three separate occasions (July, November, and March) over a 12-month period. Parsimony analyses of the molecular data for adults and progeny collected from mussels during July, November, and March revealed distinct groupings, that for the most part, corresponded to mites collected from each of the sampling periods. Many of the genetic markers obtained for male and female U. foili were not evident among the larvae or eggs, suggesting that adults obtained from a host mussel at the time of collection were not the parents of a majority of the progeny. However, female mites and eggs collected from mussels during March and November shared more markers than did females and progeny examined during July. Furthermore, many offspring in the July sampling period were found to have one or more parents absent from the sampled population. Overall, RAPD profiling appears to have limited usage in determining kinship within populations of U. foili, due to its recruitment patterns, and the relatively large number of adults and progeny per mussel. It may, however, prove to be a useful method for assessing genetic relatedness among unionicolid mussel-mites that have substantially lower population densities.

Host sex preferences and transmission success by the water mite Unionicola foili (Acari: Unionicolidae) parasitic on the midge Chironomus tentans (Diptera: Chironomidae).

This study examined whether ecoparasitic larval Unionicola foili exhibited a sex bias when infecting laboratory populations of the host insect Chironomus tentans and whether an association with male or female midges increased the likelihood of larval mites returning to the aquatic habitat. When laboratory populations of C. tentans were exposed to larval U. foili, there was a higher prevalence of mites among female hosts at emergence (17 of 30 males vs. 25 of 30 females infected by mites). However, there was no significant difference in the distribution or abundance of larvae among infected male (mean = 2.3 larvae per host) and female (mean = 2.6 larvae per host) midges. Larval mites parasitizing both male and female chironomids were more likely to return to water than could be expected by chance. Mite larvae infesting female C. tentans were more likely to return to water when female hosts deposited egg masses in water, suggesting that oviposition plays an important role in cueing larvae parasitizing female midges to detach. The mechanism responsible for increasing the likelihood that mites parasitizing male hosts return to water remains unclear. Future studies will address the possibility of parasite-mediated changes in host behavior.