scholarly journals Diversity, Host Ranges, and Potential Drivers of Speciation Among the Inquiline Enemies of Oak Gall Wasps (Hymenoptera: Cynipidae)

2020 ◽  
Vol 4 (6) ◽  
Author(s):  
Anna K G Ward ◽  
Sofia I Sheikh ◽  
Andrew A Forbes

Abstract Animals that exploit living spaces of other animals (inquilines) may have specialized traits that adapt them to extended phenotypes of their ‘hosts’. These adaptations to host traits may incur fitness trade-offs that restrict the host range of an inquiline such that shifts to new hosts might trigger inquiline diversification. Speciation via host shifting has been studied in many animal parasites, but we know less about the role of host shifts in inquiline speciation. Synergus Hartig (Hymenoptera: Cynipidae: Synergini) is a speciose but taxonomically challenging genus of inquilines that feed inside galls induced by oak gall wasps (Hymenoptera: Cynipidae: Cynipini). 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 complexes of two to five species, while three species fit no prior description. We find evidence that oak tree phylogeny and host gall morphology define axes of specialization for Synergus. The North American Synergus have experienced several transitions among gall hosts and tree habitats and their host use is correlated with reproductive isolation. It remains too early to tell whether shifts to new hosts initiate speciation events in Synergus inquilines of oak gall wasps, or if host shifts occur after reproductive isolation has already evolved.

Author(s):  
Anna KG Ward ◽  
Sofia I Sheikh ◽  
Andrew A Forbes

ABSTRACTAnimals 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.


2012 ◽  
Vol 2012 ◽  
pp. 1-8 ◽  
Author(s):  
Scott P. Egan ◽  
Glen R. Hood ◽  
James R. Ott

Habitat isolation occurs when habitat preferences lower the probability of mating between individuals associated with differing habitats. While a potential barrier to gene flow during ecological speciation, the effect of habitat isolation on reproductive isolation has rarely been directly tested. Herein, we first estimated habitat preference for each of six populations of the gall waspBelonocnema treataeinhabiting eitherQuercus virginianaorQ. geminata. We then estimated the importance of habitat isolation in generating reproductive isolation betweenB. treataepopulations that were host specific to eitherQ. virginianaorQ. geminataby measuring mate preference in the presence and absence of the respective host plants. All populations exhibited host preference for their native plant, and assortative mating increased significantly in the presence of the respective host plants. This host-plant-mediated assortative mating demonstrates that habitat isolation likely plays an important role in promoting reproductive isolation among populations of this host-specific gall former.


Author(s):  
Kelly L. Weinersmith ◽  
Andrew A. Forbes ◽  
Anna K.G. Ward ◽  
Pedro F. P. Brandão-Dias ◽  
Y. Miles Zhang ◽  
...  

AbstractCynipid 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.


2020 ◽  
Vol 113 (5) ◽  
pp. 373-388 ◽  
Author(s):  
Kelly L Weinersmith ◽  
Andrew A Forbes ◽  
Anna K G Ward ◽  
Pedro F P Brandão-Dias ◽  
Y Miles Zhang ◽  
...  

Abstract Cynipid gall wasps play an important role in structuring oak arthropod 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 Quercus 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 arthropods 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 galls support a diverse arthropod community, including over 25 parasitoids, inquilines, and other associated arthropods spanning 5 orders and 16 families.


2019 ◽  
Vol 15 (9) ◽  
pp. 20190428 ◽  
Author(s):  
Anna K. G. Ward ◽  
Omar S. Khodor ◽  
Scott P. Egan ◽  
Kelly L. Weinersmith ◽  
Andrew A. Forbes

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 host's behaviour, the added complexity of that manipulation may further limit the parasite's host range. However, this is rarely tested. The recently described crypt-keeper wasp, Euderus set , changes the behaviour of the gall wasp Bassettia pallida such that B. pallida chews a significantly smaller exit hole in the side of its larval chamber and ‘plugs’ that hole with its head before dying. Euderus set benefits from this head plug, as it facilitates the escape of the parasitoid from the crypt after it completes development. Here, we find direct and indirect evidence that E. set attacks and manipulates the behaviour of at least six additional gall wasp species, and that these hosts are taxonomically diverse. Interestingly, each of E. set 's hosts has converged upon similarities in their extended phenotypes: the galls they induce on oaks share characters that may make them vulnerable to attack by E. set . The specialization required to behaviourally manipulate hosts may be less important in determining the range of hosts in this parasitoid system than other dimensions of the host–parasitoid interaction, like the host's physical defences.


2019 ◽  
Author(s):  
Anna K. G. Ward ◽  
Omar S. Khodor ◽  
Scott P. Egan ◽  
Kelly L. Weinersmith ◽  
Andrew A. Forbes

AbstractParasites 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 host’s behavior, the complexity of that manipulation may further limit the parasite’s host range. The recently described crypt-keeper wasp,Euderus set, changes the behavior of the gall waspBassettia pallidasuch thatB. pallidachews an incomplete exit hole in the side of its larval chamber and “plugs” that hole with its head.E. setbenefits 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 toBassettiahosts. We find thatE. setattacks and manipulates the behavior of at least six additional gall wasp species, and that these hosts are taxonomically diverse. Interestingly, each ofE. set’s hosts has converged upon similarities in their extended phenotypes: the galls they induce on oaks share characters that may make them vulnerable to attack byE. 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 host’s physical defenses.


2021 ◽  
Vol 12 ◽  
Author(s):  
Dao-Hong Zhu ◽  
Cheng-Yuan Su ◽  
Xiao-Hui Yang ◽  
Yoshihisa Abe

The phage WO was characterized in Wolbachia, a strictly intracellular bacterium causing several reproductive alterations in its arthropod hosts. This study aimed to screen the presence of Wolbachia and phage WO in 15 gall wasp species from six provinces of southern China to investigate their diversity and prevalence patterns. A high incidence of Wolbachia infection was determined in the gall wasp species, with an infection rate of 86.7% (13/15). Moreover, seven species had double or multiple infections. All Wolbachia-infected gall wasp species were found to harbor phage WO. The gall wasp species infected with a single Wolbachia strain were found to harbor a single phage WO type. On the contrary, almost all species with double or multiple Wolbachia infections harbored a high level of phage WO diversity (ranging from three to 27 types). Six horizontal transfer events of phage WO in Wolbachia were found to be associated with gall wasps, which shared identical orf7 sequences among their respective accomplices. The transfer potentially took place through gall inducers and associated inquilines infected with or without Wolbachia. Furthermore, 10 putative recombination events were identified from Andricus hakonensis and Andricus sp2, which harbored multiple phage WO types, suggesting that intragenic recombination was the important evolutionary force, which effectively promoted the high level of phage WO diversity associated with gall wasps.


2002 ◽  
Vol 24 (1) ◽  
pp. 1-20 ◽  
Author(s):  
ANDREW BAINES

In reading archaeological texts, we expect to be engaged in a characteristically archaeological discourse, with a specific and recognisable structure and vocabulary. In evaluating the published work of 19th Century antiquarians, we will inevitably look for points of contact between their academic language and our own; success or failure in the identification of such points of contact may prompt us to recognise a nascent archaeology in some writings, while dismissing others as naïve or absurd. With this point in mind, this paper discusses the written and material legacies of three 19th Century antiquarians in the north of Scotland who worked on a particular monument type, the broch. The paper explores the degree to which each has been admitted as an influence on the development of the broch as a type. It then proceeds to compare this established typology with the author's experiences, in the field, of the sites it describes. In doing so, the paper addresses wider issues concerning the role of earlier forms of archaeological discourse in the development of present day archaeological classifications of, and of the problems of reconciling such classifications with our experiences of material culture.


2020 ◽  
Vol 58 ◽  
pp. 181-192
Author(s):  
Anna A. Komzolova

One of the results of the educational reform of the 1860s was the formation of the regular personnel of village teachers. In Vilna educational district the goal was not to invite teachers from central Russia, but to train them on the spot by establishing special seminaries. Trained teachers were supposed to perform the role of «cultural brokers» – the intermediaries between local peasants and the outside world, between the culture of Russian intelligentsia and the culture of the Belarusian people. The article examines how officials and teachers of Vilna educational district saw the role of rural teachers as «cultural brokers» in the context of the linguistic and cultural diversity of the North-Western Provinces. According to them, the graduates of the pedagogical seminaries had to remain within the peasant estate and to keep in touch with their folk «roots». The special «mission» of the village teachers was in promoting the ideas of «Russian elements» and historical proximity to Russia among Belarusian peasants.


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