Biology subjects

Forbes, A. A.

Publications and source records attributed to Forbes, A. A..

16 recordsLinked to original sources

bioRxiv2026

Assessing positive selection in centromere-associated kinetochore proteins across Metazoan groups.

Centromeres are comprised of long stretches of repetitive DNA that evolve rapidly in organisms across the tree of life. Consistent selfish centromere evolution can also have cascading effects - driving rapid evolution in interacting kinetochore proteins - possibly to maintain centromere-kinetochore compatibility. Effects of selfishly evolving centromeres on interacting proteins are most heavily studied in the inner kinetochore and assembly proteins including the constitutive centromere-associated network proteins CENP-A and CENP-C with some exploration of the extended effects to other kinetochore-associated protein complexes. While rapid evolution of the centromere has been broadly studied in many organisms, studies assessing positive selection in centromere-associated kinetochore proteins have largely focused on Drosophila. Here, we tested the hypothesis that signatures of positive selection would be present in outer kinetochore and condensin genes in diverse animal groups. We selected two protein complexes -the Condensin I complex and the Mis12 Complex - to test for positive selection in parasitic wasps, two groups of ray-finned fishes (including the amazon molly an asexual diploid exempt from centromere drive), and two groups of primates. We did not find selection using any test in any protein in the amazon molly but did find sporadic positive selection in proteins in both complexes across all groups.

evolutionary biology↗

bioRxiv2025

Periodical cicadas suffer legacy effects of century-old forest removal

Periodical cicada (Magicicada spp.) ranges have continued to decline even as their forest habitat has rebounded from historic lows. This counterintuitive trajectory may be a consequence of Magicicadas limited dispersal capabilities combined with their requirement that individual host trees survive across decades. We studied the relationship between >150 years of landscape change and brood XIII periodical cicada activity in Johnson County, IA. We found that the presence of Magicicada breeding populations was better predicted by tree cover in 1930 than by present-day tree cover, with cicadas primarily relegated to sites that did not fall below 17.4% tree cover. While total forest has increased since 1930, this is a net increase, inclusive of considerable turnover every 17 years, and cicada ranges have declined more than they have expanded. A model based on empirical measures of forest turnover and cicada dispersal distances suggests that cicada ranges will continue to decline under nearly all conditions. A managed relocation of 20,126 adult Magicicada to a restored forest failed, reinforcing that dispersal to new sites cannot be artificially accelerated to offset declines. Preservation of forest habitat across centuries may be the only bollard standing between periodical cicadas and their extinction.

ecology↗

bioRxiv2023

Speciation in kleptoparasites of oak gall wasps often correlates with shifts into new tree habitats, tree organs, and gall morphospace

Host shifts to new plants can drive speciation for plant-feeding insects, but how commonly do host shifts also drive diversification for the parasites of those same insects? Oak gall wasps induce galls on oak trees and shifts to novel tree hosts and new tree organs have been implicated as drivers of oak gall wasp speciation. Gall wasps are themselves attacked by many insect parasites, which must find their hosts on the correct tree species and organ, but which also must navigate the morphologically variable galls with which they interact. Thus, we ask whether host shifts to new trees, organs, or gall morphologies correlate with gall parasite diversification. We delimit species and infer phylogenies for two genera of gall kleptoparasites, Synergus and Ceroptres, reared from a variety of North American oak galls. We find that most species were reared from galls induced by just one gall wasp species, and no parasite species was reared from galls of more than four species. Most kleptoparasite divergence events correlate with shifts to non-ancestral galls. These shifts often involved changes in tree habitat, gall location, and gall morphology. Host shifts are thus implicated in driving diversification for both oak gall wasps and their kleptoparasitic associates.

evolutionary biology↗

bioRxiv2022

Host-enemy interactions provide limited biotic resistance for a range-expanding species via reduced apparent competition

As species ranges shift in response to anthropogenic change, they lose coevolved or coadapted interactions and gain novel ones in recipient communities. Range-expanding species may lose or experience weak antagonistic interactions with competitors and enemies, and traits of interacting species will determine the strength of interactions. We leveraged a poleward range expansion of an oak gall wasp that co-occurs on its host plant with other gall wasp species and interacts with shared natural enemies (largely parasitoid wasps). We created quantitative host-parasitoid interaction networks by sampling galls on 400 trees. We compared network structure and function and traits of hosts and parasitoids in the native and expanded range. Interaction networks were less diverse in the expanded range, with low complementarity of parasitoid assemblages among hosts. While whole networks were more generalized in the expanded range, interactions with the range-expanding species were more specialized. This was not due to a loss of specialist enemies but weak apparent competition by shared generalist enemies. Phenological divergence of enemy assemblages attacking the novel and co-occurring hosts was greater in the expanded range that may contribute to weak apparent competition. Given the rate and extent of anthropogenic-driven range expansions, it is pressing to uncover how complex biotic interactions are reassembled.

ecology↗

bioRxiv2022

Stronger sexual dimorphism in fruit flies may be favored when congeners are present and females actively search for mates

Why are some species sexually dimorphic while other closely related species are not? When the degree of sexual dimorphism varies within a genus, an integrative phylogenetic approach may help reveal underlying patterns favoring the evolution of dimorphism. While all female flies in genus Strauzia - a genus of true fruit flies - share a multiply-banded wing pattern, males of four species have patterns wherein bands have "coalesced" into a continuous dark streak across much of the wing. We find that the origin of coalesced male wing patterns and pronounced differences in male wing shape correlate with the presumed origin of host plant sharing in this genus. A survey of North American Tephritidae finds just three other genera with specialist species that share host plants. Each has one or more congeners with wing patterns unusual for that genus, and just one genus, Eutreta, has those unusual wing patterns only in the male sex. Eutreta is also the only other genus among this subset wherein, like Strauzia, males hold territories while females search for mates. Sharing the same hosts may result in reproductive character displacement, and when coupled with a biology wherein females actively search for males, may specifically favor sexually dimorphic wing patterns.

evolutionary biology↗

bioRxiv2022

The arthropod associates of 155 North American cynipid oak galls

The identities of most arthropod associates of cynipid-induced oak galls in the western Palearctic are generally known. However, a comprehensive accounting of associates has been performed for only a small number of the galls induced by the estimated 700 species of cynipid gall wasp in the Nearctic. This gap in knowledge stymies many potential studies of diversity, coevolution, and community ecology, for which oak gall systems are otherwise ideal models. We report rearing records of insects and other arthropods from more than 527,306 individual galls representing 201 different oak gall types collected from 32 oak tree species in North America. Of the 201 gall types collected, 155 produced one or more animals. A total of 151,075 animals were found in association with these 155 gall types, and of these 61,044 (40.4%) were gall wasps while 90,031 (59.6%) were other arthropods. We identified all animals to superfamily, family, or, where possible, to genus. We provide raw numbers and summaries of collections, alongside notes on natural history, ecology, and previously published associations for each taxon. For eight common gall-associated genera (Synergus, Ceroptres, Euceroptres, Ormyrus, Torymus, Eurytoma, Sycophila, and Euderus), we also connect rearing records to gall wasp phylogeny, geography, and ecology - including host tree and gall location (host organ), and their co-occurrence with other insect genera. Though the diversity of gall wasps and the large size of these communities is such that many Nearctic oak gall-associated insects still remain undescribed, this large collection and identification effort should facilitate the testing of new and varied ecological and evolutionary hypotheses in Nearctic oak galls.

zoology↗

bioRxiv2022

Speciation in Nearctic oak gall wasps is frequently correlated with changes in host plant, host organ, or both.

Quantifying the frequency of shifts to new host plants within diverse clades of specialist herbivorous insects is critically important to understand whether and how host shifts contribute to the origin of species. Oak gall wasps (Hymenoptera: Cynipidae: Cynipini) comprise a tribe of ~1000 species of phytophagous insects that induce gall formation on various organs of trees in the family Fagacae, -- primarily the oaks (genus Quercus; ~435 sp). The association of oak gall wasps with oaks is ancient (~50 my), and most oak species are galled by one or more gall wasp species. Despite the diversity of both gall wasp species and their plant associations, previous phylogenetic work has not identified a strong signal of host plant shifting among oak gall wasps. However, most emphasis has been on the Western Palearctic and not the Nearctic where both oaks and oak gall wasps are considerably more species rich and where oaks are more phylogenetically diverse. We collected 86 species of Nearctic oak gall wasps from 10 of the 14 major clades of Nearctic oaks and sequenced >1000 Ultra Conserved Elements (UCEs) and flanking sequences to infer wasp phylogenies. We assessed the relationships of Nearctic gall wasps to one another and, by leveraging previously published UCE data, to the Palearctic fauna. We then used phylogenies to infer historical patterns of shifts among host tree species and tree organs. Our results indicate that oak gall wasps have moved between the Palearctic and Nearctic at least four times, that some Palearctic clades have their proximate origin in the Nearctic, and that gall wasps have shifted within and between oak tree sections, subsections, and organs considerably more often than the analysis of previous data have suggested. Given that host shifts have been demonstrated to drive reproductive isolation between host-associated populations in other phytophagous insects, our analyses of Nearctic gall wasps suggest that host shifts are key drivers of speciation in this clade, especially in hotspots of oak diversity. Though formal assessment of this hypothesis requires further study, two putatively oligophagous gall wasp species in our dataset show signals of host-associated genetic differentiation unconfounded by geographic distance, suggestive of barriers to gene flow associated with the use of alternative host plants.

evolutionary biology↗

bioRxiv2022

Delimiting the cryptic diversity and host preferences of Sycophila parasitoid wasps associated with oak galls using phylogenomic data

Cryptic species diversity is a major challenge for the species-rich community of parasitoids attacking oak gall wasps due to a high degree of sexual dimorphism, morphological plasticity, small size, and poorly known biology. As such, we know very little about the number of species present, nor the evolutionary forces responsible for generating this diversity. One hypothesis is that trait diversity in the gall wasps, including the morphology of the galls they induce, has evolved in response to selection imposed by the parasitoid community, with reciprocal selection driving diversification of the parasitoids. Using a rare, continental-scale data set of Sycophila parasitoid wasps reared from 44 species of cynipid galls from 18 species of oak across the US, we combined mitochondrial DNA barcodes, Ultraconserved Elements (UCEs), morphological, and natural history data to delimit putative species. Using these results, we generate the first large-scale assessment of ecological specialization and host association in this species-rich group, with implications for evolutionary ecology and biocontrol. We find most Sycophila target specific subsets of available cynipid host galls with similar morphologies, and generally attack larger galls. Our results suggest that parasitoid wasps such as Sycophila have adaptations allowing them to exploit particular host trait combinations, while hosts with contrasting traits are resistant to attack. These findings support the tritrophic niche concept for the structuring of plant-herbivore-parasitoid communities.

evolutionary biology↗

bioRxiv2021

Ormyrus labotus Walker (Hymenoptera: Ormyridae): another generalist that should not be a generalist is not a generalist

Several recent reappraisals of supposed generalist parasite species have revealed hidden complexes of species, each with considerably narrower host ranges. Parasitic wasps that attack gall-forming insects on plants have life history strategies that are thought to promote specialization, and though many species are indeed highly specialized, others have been described as generalist parasites. Ormyrus labotus Walker (Hymenoptera: Ormyridae) is one such apparent generalist, with rearing records spanning more than 65 host galls associated with a diverse set of oak tree species and plant tissues. We pair a molecular approach with morphology, host ecology, and phenological data from across a wide geographic sample to test the hypothesis that this supposed generalist is actually a complex of several more specialized species. We find 16-18 putative species within the morphological species O. labotus, each reared from only 1-6 host gall types, though we identify no single unifying axis of specialization. We also find cryptic habitat specialists within two other named Ormyrus species. Our study suggests that caution should be applied when considering host ranges of parasitic insects described solely by morphological traits, particularly given their importance as biocontrol organisms and their role in biodiversity and evolutionary studies.

evolutionary biology↗

bioRxiv2020

Genome evolution in a putatively asexual wasp

Asexual lineages are destined for extinction--a result predicted by theory and revealed in practice. Short-term benefits of asexuality are eventually outstripped by their fitness costs: losses of sex and recombination are together expected to reduce efficacy of selection, increase mutation load, and thus, lower fitness. We characterized genomic patterns of accumulating mutations in Diachasma muliebre, a parasitic wasp that has apparently lost sex, an inference supported by many decades of field collections of 1000s of individuals in which only females were found. The split between D. muliebre and its closest sexual relative, Diachasma ferrugineum, is quite recent, allowing us to observe initial events in the evolution of this putative asexual species. First, we find a faster rate of molecular evolution across the D. muliebre genome. In addition, we observed a marked excess of replacement nucleotide substitutions in orthologous genes in the putatively asexual D. muliebre when compared to D. ferrugineum. This pattern directly indicates genome-wide relaxed selection in this young, putatively asexual species, the resulting mutational load from which is expected to ultimately lead to extinction. However, these genomic effects occur in the presence of genomic recombination initially detected by a previous study and also supported by analyses of genome-wide substitution rates within codons. In addition, following completion of the genome sequence and its analysis, we discovered two D. muliebre males, suggesting the possibility of rare sex in this species. Haplodiploid animals, including the sexual ancestors of D. muliebre, bear small genetic loads, likely making their initial transitions to asexuality relatively benign. Paradoxically, an elevated rate of mutation accumulation resulting from asexuality, when accompanied by retention of recombination and/or rare sex, could actually be beneficial: we hypothesize that the novel variation introduced by mutation along with limited shuffling of genes may facilitate initial adaptation and extend persistence of such lineages.

evolutionary biology↗

bioRxiv2020

Diversity, host ranges, and potential drivers of speciation among the inquiline enemies of oak gall wasps.

Animals that exploit an extended phenotype (e.g., residences, resources, etc.) of other animals are called inquilines. Not strictly parasites, inquilines may nevertheless possess specialized traits that adapt them to particular dimensions of the extended phenotype of their "host". These adaptations to host traits can in turn lead to fitness tradeoffs that restrict the host range of an inquiline such that shifts to novel hosts might trigger inquiline diversification. Speciation via host shifting has been studied in many animal parasites, but we know far less about the role of host shifts in inquiline speciation. Synergus (Hymenoptera: Cynipidae: Synergini) is a speciose but taxonomically-challenging group of inquilines that feed on the tissue of galls induced by oak gall wasps (Hymenoptera: Cynipidae: Cynipini). Currently too little is known about Nearctic Synergus diversity or host associations to evaluate whether and how host use affects their diversification. Here, we report on a large collection of Synergus reared from galls of 33 oak gall wasp species in the upper Midwestern United States. We integrated DNA barcodes, morphology, ecology, and phenology to delimit putative species of Synergus and describe their host ranges. We find evidence of at least 23 Synergus species associated with the 33 gall wasp hosts. At least five previously described Synergus species are each a complex of two to five species, while three species fit no prior description. We also find strong evidence that oak tree section and host gall morphology define axes of specialization for Synergus. Without over-interpreting our singlegene tree, it is clear that the North American Synergus have experienced several transitions among gall hosts and tree habitats and that host-use is correlated with reproductive isolation, though it remains too early to tell whether shifts to new hosts are the initiators of speciation events in Synergus inquilines of oak gall wasps, or if host shifts occur after reproductive isolation has already evolved.

evolutionary biology↗

bioRxiv2020

Host shifting and host sharing in a genus of specialist flies diversifying alongside their sunflower hosts

Congeneric parasites are unlikely to specialize on the same tissues of the same host species, likely because of strong multifarious selection against niche overlap. Exceptions where multiple congeneric species overlap on the same tissues may therefore reveal important insights into the ecological factors underlying the origins and maintenance of diversity. Larvae of sunflower maggot flies in genus Strauzia feed on the pith of plants in the family Asteraceae. Although Strauzia tend to be host specialists, some species overlap in their host use. To resolve the origins of host sharing among these specialist flies, we used reduced representation genomic sequencing to infer the first multi-locus phylogeny of genus Strauzia. Our results show that Helianthus tuberosus and Helianthus grosseserratus each host three different fly species, and that the flies co-occurring on a host are not one anothers closest relatives. Though this pattern implies that host sharing is most likely the result of host shifts, these may not be host shifts in the conventional sense of an insect moving onto an entirely new plant. Many hosts of Strauzia belong to a young (1-2 MYA) clade of perennial sunflowers noted for their frequent introgression and hybrid speciation events. In at least one case, flies may have converged upon a host after their respective ancestral host plants hybridized to form a new sunflower species (H. tuberosus). Broadly, we suggest that rapid and recent adaptive introgression and speciation in this group of plants may have instigated the diversification of their phytophagous fly associates, including the convergence of >1 species onto the same shared host plants.

evolutionary biology↗

bioRxiv2020

Invertebrate Community Associated with the Asexual Generation of Bassettia pallida Ashmead (Hymenoptera: Cynipidae)

Cynipid gall wasps play an important role in structuring oak invertebrate communities. Wasps in the Cynipini tribe typically lay their eggs in oaks (Quercus L.), and induce the formation of a "gall", which is a tumor-like growth of plant material that surrounds the developing wasp. As the wasp develops, the cynipid and its gall are attacked by a diverse community of natural enemies, including parasitoids, hyperparasitoids, and inquilines. Determining what structures these species-rich natural enemy communities across cynipid gall wasp species is a major question in gall wasp biology. Additionally, gall wasps are ecosystem engineers, as the abandoned gall is used by other invertebrates. The gall-associated insect communities residing on live oaks (Quercus geminata Small and Q. virginiana Mill.) are emerging as a model system for answering ecological and evolutionary questions ranging from community ecology to the evolution of new species. Documenting the invertebrates associated with cynipids in this system will expand our understanding of the mechanisms influencing eco-evolutionary processes, record underexplored axes of biodiversity, and facilitate future work. Here, we present the community of natural enemies and other associates of the asexual generation of the crypt gall wasp, Bassettia pallida Ashmead. We compare the composition of this community to communities recently documented from two other cynipid gall wasps specializing on live oaks along the U.S. Gulf coast, Disholcaspis quercusvirens Ashmead and Belonocnema treatae Mayr. B. pallida and their crypts support a diverse arthropod community, including over 25 parasitoids, inquilines, and other associated invertebrates spanning 5 orders and 16 families.

ecology↗

bioRxiv2019

Genome of the parasitoid wasp Diachasma alloeum, an emerging model for ecological speciation and transitions to asexual reproduction

Parasitoid wasps are among the most speciose animals, yet have relatively few available genomic resources. We report a draft genome assembly of the wasp Diachasma alloeum (Hymenoptera: Braconidae), a host-specific parasitoid of the apple maggot fly Rhagoletis pomonella (Diptera: Tephritidae) and a developing model for understanding how ecological speciation can "cascade" across trophic levels. Identification of gene content confirmed the overall quality of the draft genome, and we manually annotated [~]400 genes as part of this study, including those involved in oxidative phosphorylation, chemosensation, and reproduction. Through comparisons to model hymenopterans such as the European honeybee Apis mellifera and parasitoid wasp Nasonia vitripennis, as well as a more closely related braconid parasitoid Microplitis demolitor, we identified a proliferation of transposable elements in the genome, an expansion of chemosensory genes in D. alloeum and other parasitoid wasps, and the maintenance of several key genes with known roles in sexual reproduction and sex determination. The D. alloeum genome will provide a valuable resource for comparative genomics studies in Hymenoptera as well as specific investigations into the genomic changes associated with ecological speciation and transitions to asexuality.

genomics↗

bioRxiv2019

A keeper of many crypts: the parasitoid Euderus set manipulates the behavior of a taxonomically diverse array of oak gall wasp species

Parasites of animals and plants can encounter trade-offs between their specificity to any single host and their fitness on alternative hosts. For parasites that manipulate their hosts behavior, the complexity of that manipulation may further limit the parasites host species range. The recently described crypt-keeper wasp, Euderus set, changes the behavior of the gall wasp Bassettia pallida such that B. pallida chews an incomplete exit hole in the side of its larval chamber and \"plugs\" that hole with its head. E. set benefits from this head plug, as it facilitates the escape of the parasitoid from the crypt after it completes development. Here, we ask whether this behavioral manipulator is limited to Bassettia hosts. We find that E. set attacks and manipulates the behavior of at least six additional gall wasp species, and that these hosts are taxonomically diverse. Interestingly, each of E. sets hosts has converged upon similarities in the extended phenotypes they induce in their plant hosts: the galls they induce on oaks share characters that may make them vulnerable to attack by E. set. Behavioral manipulation in this parasitoid system may be less important to its host range than other dimensions of the host-parasitoid interaction, like the hosts physical defenses.

ecology↗

bioRxiv2018

Quantifying the unquantifiable: why Hymenoptera -- not Coleoptera -- is the most speciose animal order

BackgroundWe challenge the oft-repeated claim that the beetles (Coleoptera) are the most species-rich order of animals. Instead, we assert that another order of insects, the Hymenoptera, are more speciose, due in large part to the massively diverse but relatively poorly known parasitoid wasps. The idea that the beetles have more species than other orders is primarily based on their respective collection histories and the relative availability of taxonomic resources, which both disfavor parasitoid wasps. Though it is unreasonable to directly compare numbers of described species in each order, the ecology of parasitic wasps - specifically, their intimate interactions with their hosts - allows for estimation of relative richness. We present a simple logical model that shows how the specialization of many parasitic wasps on their hosts suggests few scenarios in which there would be more beetle species than parasitic wasp species. We couple this model with an accounting of what we call the \"genus-specific parasitoid-host ratio\" from four well-studied genera of insect hosts, a metric by which to generate extremely conservative estimates of the average number of parasitic wasp species attacking a given beetle or other insect host species. Synthesis of our model with data from real host systems suggests that the Hymenoptera may have 2.5 - 3.2x more species than the Coleoptera. While there are more described species of beetles than all other animals, the Hymenoptera are almost certainly the larger order.

zoology↗