Climate,host phylogeny and the connectivity of host communities govern regional parasite assembly

Aim

Identifying barriers that govern parasite community assembly and parasite invasion risk is critical to understand how shifting host ranges impact disease emergence. We studied regional variation in the phylogenetic compositions of bird species and their blood parasites (Plasmodium and Haemoproteus spp.) to identify barriers that shape parasite community assembly.

Location

Australasia and Oceania.

Methods

We used a data set of parasite infections from >10,000 host individuals sampled across 29 bioregions. Hierarchical models and matrix regressions were used to assess the relative influences of interspecies (host community connectivity and local phylogenetic distinctiveness), climate and geographic barriers on parasite local distinctiveness and composition.

Results

Parasites were more locally distinct (co‐occurred with distantly related parasites) when infecting locally distinct hosts, but less distinct (co‐occurred with closely related parasites) in areas with increased host diversity and community connectivity (a proxy for parasite dispersal potential). Turnover and the phylogenetic symmetry of parasite communities were jointly driven by host turnover, climate similarity and geographic distance.

Main conclusions

Interspecies barriers linked to host phylogeny and dispersal shape parasite assembly, perhaps by limiting parasite establishment or local diversification. Infecting hosts that co‐occur with few related species decreases a parasite's likelihood of encountering related competitors, perhaps increasing invasion potential but decreasing diversification opportunity. While climate partially constrains parasite distributions, future host range expansions that spread distinct parasites and diminish barriers to host shifting will likely be key drivers of parasite invasions.     

Host species identity in annual Brassicaceae has a limited effect on the assembly of root-endophytic fungal communities

ABSTRACT

Background

Establishing the contribution of host plant genotype to the assembly of root-associated microbial communities can be challenging, since multiple ecological factors may confound the effects purely due to the host, and plant species often do not occur sympatrically.     

Host range evaluation of the leaf-feeding beetle,Zygogramma bicolorata and the stem-boring weevil,Listronotus setosipennis demonstrates their suitability for biological control of the invasive weed,Parthenium hysterophorus in Ethiopia

The invasive weed, parthenium (Parthenium hysterophorus L.) (Asteraceae: tribe Heliantheae), damages agriculture, adversely impacts biodiversity and is hazardous to human and animal health in Ethiopia. The host range of two natural enemies, a leaf-feeding beetle, Zygogramma bicolorata (Coleoptera: Chrysomelidae) and a stem-boring weevil, Listronotus setosipennis (Coleoptera: Curculionidae) was evaluated for biological control of the weed in Ethiopia. The specificity of Z. bicolorata and L. setosipennis was assessed against 29 and 31 non-target plant species, respectively. The host range of Z. bicolorata and L. setosipennis was first assessed using no-choice tests to examine their oviposition and feeding response on non-target plants. Although oviposition by Z. bicolorata occurred on six non-target species in four Asteraceae species in no-choice tests, it was significantly lower than on parthenium and no larvae developed. Zygogramma bicolorata nibbled the leaves of one of the five niger seed (Guizotia abyssinica L. – an oil seed crop closely related to parthenium) cultivars tested, but feeding and oviposition were significantly less than on parthenium. Furthermore, choice tests indicated that Z. bicolorata did not oviposit nor feed on G. abyssinica when parthenium was present. In no-choice tests, L. setosipennis did not oviposit on any of the non-target species assessed. Mean oviposition on parthenium was 39.0?±?3.4 eggs per plant whereas no eggs were laid on any of the 31 species tested. Based on these and other host range tests, permission was obtained to field release Z. bicolorata and L. setosipennis in Ethiopia.     

Getting there and around: Host range oscillations during colonization of the Canary Islands by the parasitic nematode Spauligodon

Episodes of expansion and isolation in geographic range over space and time, during which parasites have the opportunity to expand their host range, are linked to the development of host–parasite mosaic assemblages and parasite diversification. In this study, we investigated whether island colonization events lead to host range oscillations in a taxon of host‐specific parasitic nematodes of the genus Spauligodon in the Canary Islands. We further investigated whether range oscillations also resulted in shifts in host breadth (i.e., specialization), as expected for parasites on islands. Parasite phylogeny and divergence time estimates were inferred from molecular data with Bayesian methods. Host divergence times were set as calibration priors after a priori evaluation with a global‐fit method of which individual host–parasite associations likely represent cospeciation links. Parasite colonization history was reconstructed, followed by an estimation of oscillation events and specificity level. The results indicate the presence of four Spauligodon clades in the Canary Islands, which originated from at least three different colonization events. We found evidence of host range oscillations to truly novel hosts, which in one case led to higher diversification. Contemporary host–parasite associations show strong host specificity, suggesting that changes in host breadth were limited to the shift period. Lineages with more frequent and wider taxonomic host range oscillations prior to the initial colonization event showed wider range oscillations during colonization and diversification within the archipelago. Our results suggest that a lineage's evolutionary past may be the best indicator of a parasite's potential for future range expansions.     

Comparison of lepidopteran larval communities among tree species in a temperate deciduous forest, Japan

Abstract.  1. The enormous diversity of phytophagous insects in forest canopies is hypothesised to be supported by the number of herbivorous species per host tree species or host specificity. It is therefore necessary to examine the effect of host plant species on compositional changes in the herbivore communities.
2. The lepidopteran larval communities were examined in the canopies of 10 tree species in a temperate deciduous forest of Japan. The phylogeny and leaf flush phenology of host plant species were taken into account as factors affecting the herbivore community assembly.
3. Examination of seasonal changes in the larval community structures on each tree species showed that larval species richness, abundance, and evenness decreased significantly from spring to summer. Larval species richness and abundance were characterised by family-level phylogenetic differences among tree species, whereas evenness was determined at a higher taxonomic level.
4. Compositional changes in the larval communities among tree species showed a remarkable pattern, with a phylogenetic effect at a high taxonomic level in spring, similar to evenness, but a phenological effect in summer. This suggests that host specificity could support the lepidopteran larval diversity in spring.
5. These results suggest that the differences in host utilisation of the herbivore, which reflects the phylogenetic effect of the host plants, can be important as a factor affecting the diversity of lepidopteran larval communities in temperate forests.     

Host–parasite relations of an angiospermous root parasite (Thonningia sanguinea Vahl) in logged and unlogged sites of Budongo forest reserve, western Uganda

Host–parasite relationships of an angiospermous root parasite ( Thonningia sanguinea ) were investigated in logged and unlogged sites of Budongo Forest Reserve. Host trees were identified and their diameters measured in 20 × 20 m plots established randomly in sites where the parasites were presumed to occur. The distance of the point of attachment of the parasite from the base of the host stem was determined and overstorey density measured at the centre of each plot. There were more parasites in the logged than in the unlogged sites (878 and 425 individual parasites ha⁻¹ respectively). The parasite was not host specific but Alchornea laxiflora (Benth) Pax and K.Hoffm, Celtis mildbraedii Engl and Lasiodiscus mildbraedii Engl had relatively more parasites than other species. The parasite could be found within a radius of 2 m from the base of the host stem. The diameter of hosts ranged from 1 to 95 cm. There was a positive correlation between overstorey density and occurrence of T. sanguinea . Conservation of T. sanguinea, therefore, requires maintenance of intact forests with closed canopies rather than logged sites with many gaps and hence low overstorey density.     

Biological control of marine invasive species: cautionary tales and land-based lessons

Biological control (biocontrol) has successfully regulated pest populations in terrestrial agroecosystems, but it has also caused negative unintended consequences for native species. Marine biologists and resource managers have recently published a growing number of proposals to include biocontrol in integrated pest management programs in oceans, seas and estuaries. Here, I review six ecologically and taxonomically diverse case studies of marine biocontrol programs at various stages of planning and implementation. Proposals include viral or microbial control of harmful algal blooms, predatory control of the ctenophore Mnemiopsis leidyi in the Black Sea, parasitic regulation of the European green crab Carcinus maenas, castration by ciliates of the seastar Asterias amurensis in Australia, herbivory of the toxic green alga Caulerpa taxifolia in the Mediterranean by sacoglossan sea slugs, and insect biocontrol by the planthopper Prokelesia marginata to ameliorate ecological impacts of the saltmarsh cordgrass Spartina alterniflora. Where data exist, I evaluate these examples in terms of lessons marine invasion biologists can glean from the rich history of terrestrial biocontrol, and explicitly contrast agroecosystems with invaded marine habitats. Host specificity cannot be guaranteed in the marine biocontrol proposals examined. Feasible alternatives to classical biocontrol in the marine realm should be emphasized, including more investment in invasion prevention tools, early detection and eradication while invasions are small, and increased attention to native natural enemies to control exotic pests. Biocontrol in marine habitats is risky: it poses many more uncertainties and has a much sparser history than its counterpart on land.     

Investigations on the Host Specificity of Dihomoxenous Sarcosporidia in the Intermediate and Definitive Host

ABSTRACT. Cross-transmission experiments were performed in order to determine the host specificity in the intermediate and definitive hosts of the four described dihomoxenous Sarcocystis species, S. gallotiae, S. stehlinii, S. simonyi , and S. dugesii from lacertid lizards of the genera Gallotia and Podarcis from the Macaronesian Islands. Sarcocysts of either species from experimentally infected lizards were fed to a variety of laboratory-bred lizard species of the genera Gallotia, Lacerta , and Podarcis . These sarcocysts proved to be infectious to all examined animals, showing no definitive host specificity in the tested genera. Lizards of the genera Chalcides and Tarentola , however, were not susceptible definitive hosts for S. gallotiae . The inoculation of experimentally obtained sporocysts of each of the four Sarcocystis species to various lacertid lizard species revealed varying degrees of intermediate host specificity, generally demonstrating each native host to be the most susceptible.     

Host specificity of parasitoids (Encyrtidae) toward armored scale insects (Diaspididae): Untangling the effect of cryptic species on quantitative food webs

Host specificity of parasitoids may be measured by various specialization indices to assess the variation of interaction strength among species and the structure of the wider interaction network. However, the conclusions from analyses at the species and network levels may differ, which remains poorly explored. In addition, the recovery of cryptic species of hosts and parasitoids with molecular data may affect the structure of inferred interaction links. We quantified host specificity of hymenopteran parasitoids (family Encyrtidae) on armored scale insects (Hemiptera: Diaspididae) from a wide geographic sampling range across the Chinese Mainland based on both morphological and molecular species delimitation. Mitochondrial COI and nuclear 28S markers detected high cryptic species diversity in the encyrtids and to a lesser degree in the diaspidids, which divided generalist morphospecies into complexes of specialists and generalists. One‐to‐one reciprocal host–parasite links were increased in the molecular data set, but different quantitative species‐level indices produced contrasting estimates of specificity from various one‐to‐multiple and multiple‐to‐multiple host–parasite links. Network indices calculated from DNA‐based species, compared to morphology‐based species definitions, showed lower connectance and generality, but greater specialization and compartmentalization of the interaction network. We conclude that a high degree of cryptic species in host–parasitoid systems refines the true network structure and may cause us overestimating the stability of these interaction webs.     

Host specificity assessment and potential impact of Megamelus scutellaris (Hemiptera: Delphacidae) on waterhyacinth Eichhornia crassipes (Pontederiales: Pontederiaceae)

The delphacid Megamelus scutellaris Berg was evaluated for host specificity and potential impact as part of a biological control program targeting Eichhornia. crassipes. Survival and development of adults and nymphs were used as metrics with no-choice, two-choice, nymph transfer, and sustainability tests conducted under quarantine conditions. A total of 69 plant species were tested including 12 from the Pontederiaceae (including E. crassipes). Additionally, 27 native and 5 exotic associated wetland species and 11 economic species were tested. Megamelus scutellaris exhibited a high level of oviposition and developmental fidelity to E. crassipes by failing to sustain populations on any non-target test plant past the F₁ generation. Nymph transfer tests which simulated potential spill-over events found that survival was virtually non-existent on associated wetland plants, regardless of taxonomic relatedness, including on Pontederia cordata, an important and widespread native species. Eichhornia crassipes plants exposed to two consecutive generations of feeding produced 66.9% less biomass and 73.4% fewer leaves than those in the controls. We conclude that Megamelus scutellaris is safe to release on E. crassipes in the United States.     

Reconciling genetic expectations from host specificity with historical population dynamics in an avian brood parasite, Horsfield's Bronze-Cuckoo Chalcites basalis of Australia

Mitochondrial DNA (mtDNA) is being used increasingly to explore the evolution of host specificity in avian brood parasites. A stable coevolutionary equilibrium between multiple phylogenetically unrelated hosts and a brood parasitic species predicts that mtDNA diversity in the parasite should be relatively deep and phylogenetically structured. Also, the different intraspecific clades resulting from parasitism to multiple sympatric hosts should themselves occur sympatrically. However, mtDNA diversity in brood parasites is as susceptible to effects of historical population dynamics as in any species. We demonstrate the relevance of these dynamics to the use of mtDNA in understanding coevolution between an Australian brood-parasite, Horsfield's Bronze-Cuckoo Chalcites basalis and its hosts, Malurus fairy-wrens and Acanthiza thornbills. Previous ecological and behavioural analyses argue that Malurus- and Acanthiza-specific host races exist in C. basalis. Yet mtDNA diversity in C. basalis is low and phylogenetically unstructured (mean sequence divergence 0.15 +/- 0.07%, range 0.00%-0.31%) and tests of mtDNA neutrality and range expansion vs. population stability (Tajima's D, Fu & Li's F* and D*, Fu's F(S), mismatch analyses) all indicate that C. basalis has expanded its range very recently, probably within the last few tens of thousands of years following climatic amelioration after a peak of aridity in the late Pleistocene. The low mtDNA diversity and its lack of phylogenetic structure in C. basalis deny the existence of evolutionarily long-term stable host races in C. basalis but not the possibility of recently evolved ones. They highlight the need for renewed behavioural and ecological study of the relationship between C. basalis and its hosts. Our findings illustrate the need to understand the evolutionary context in which a brood parasite and its hosts have evolved if mtDNA data are to be used in testing hypotheses concerning the origin and maintenance of host specificity. They also add to the growing body of work illustrating the use of mismatch analyses and Fu's F(S) in detecting range expansions.     

花曲柳窄吉丁的寄主植物范围、危害和防治对策

花曲柳窄吉丁(emerald ash borer, EAB) Agrilus planipennis Fairmaire是木犀科（Oleaceae）梣属Fraxinus植物树木的毁灭性蛀干害虫,目前发现其主要危害欧梣亚属Subgen. Fraxinus欧梣组(Section Fraxinus)和象蜡树组（Section Meliodes）的一些树木种类,如美国白蜡Fraxinus americana、洋白蜡F. pennsylvanica var subintegerrima、绒毛白蜡F. velutina、黑白蜡F. nigra和水曲柳F. mandshurica等。与早期文献上的记载不同,很少发现花曲柳窄吉丁对苦枥木亚属Subgen. Ornus白蜡树组(Section Ornaster)的白蜡树F. chinensis Roxb和花曲柳F. rhynchophylla造成危害。不同寄主白蜡树对花曲柳窄吉丁的敏感性不同,洋白蜡、绒毛白蜡等北美白蜡树种受其危害重于亚洲的水曲柳。这可能是由于长期的适应进化,分布在亚洲地区的梣属寄主树木发展了对花曲柳窄吉丁的抗虫性。中国部分地区发生的花曲柳窄吉丁灾害应该归属于引进的北美白蜡树种对本地害虫敏感所致。在天津、黑龙江、吉林、辽宁、河北、山东、四川、台湾、内蒙古和新疆等省市和自治区,都曾发现花曲柳窄吉丁的危害或分布。白蜡属树木广泛分布于我国除了青藏地区和海南省之外的大部分地区,其中大面积引种北美白蜡树种的北方和西北地区,以及近年来种植发展水曲柳的东北地区都是花曲柳窄吉丁这种毁灭性蛀干害虫的潜在发生地。花曲柳窄吉丁在我国部分地区为检疫性林业害虫,做好检疫是杜绝其扩散蔓延的首要方法。我国局部地区出现花曲柳窄吉丁灾害说明,北美白蜡树种在我国的适应性还需要进行更进一步科学的全面的风险评估,在我国生态建设中应重视利用我国本地的白蜡树种。在花曲柳窄吉丁灾害的治理中,可采用营造混交林以降低蔓延的速度和发挥自然控制能力,加强肥水管理以增强树木的抗虫能力,清除被害木消灭虫源,以及幼虫期树干注射、成虫期喷洒化学农药等防治措施。     

Suitability of four palm species for the development of the invasive pest Brontispa longissima (Coleoptera: Chrysomelidae) in the field

The coconut hispine beetle Brontispa longissima (Gestro) supposedly originated in Indonesia and Papua New Guinea. It is a serious invasive pest of the coconut palm Cocos nucifera L. in Southeast and East Asia. In Japan, it has established itself using Satakentia liukiuensis (Hatushima) H.E. Moore as a main host on Ishigaki and Iriomote Islands where C. nucifera is rare. To assess the probability of further establishment of B. longissima in novel habitats where C. nucifera and S. liukiuensis are not available, we examined the suitability of four common palm species in Japan for oviposition and immature development of B. longissima: Chrysalidocarpus lutescens (Bory) H. Wendl., Phoenix roebelenii O'Brien, S. liukiuensis and Washingtonia filifera (Linden ex André) H. Wendl. When seedlings of the four palm species were placed in pots in an experimental field on Ishigaki Island, all four species were inhabited and infested by wild B. longissima adults. Oviposition and immature development were observed on P. roebelenii and S. liukiuensis but not on C. lutescens and W. filifera. When field‐collected adults were released into mesh bags enclosing the potted seedlings, they oviposited on all four species. The eggs developed into adults on P. roebelenii, S. liukiuensis and W. filifera. On C. lutescens, however, hatched larvae died during the first or second instar.     

Host effects on ectomycorrhizal fungal communities: insight from eight host species in mixed conifer-broadleaf forests

To advance our understanding of host effects on the community structure of ectomycorrhizal fungi (EMF), EMF communities were compared among different host species, genera and families in two mixed conifer-broadleaf forests in Japan. Using molecular identification methods we examined EMF root tips of eight coexisting species belonging to six genera (three families): Abies and Tsuga (Pinaceae), Betula and Carpinus (Betulaceae) and Fagus and Quercus (Fagaceae). In total, 205 EMF species were detected, and the total richness was estimated to exceed 300 species using major estimators. Of the 55 EMF species occurring three or more times, eight showed significantly biased host preference. A Mantel test showed a significantly negative correlation between EMF community similarity and host taxonomic distance. Detrended correspondence analysis separated EMF communities mainly by host taxonomic and successional status. Thus, EMF communities are similar on hosts with similar taxonomic and successional status. A significant proportion of EMF exhibited host specificity, which may contribute to the extremely diverse EMF community in conifer-broadleaf forests.     

Host specificity of a generalist parasite: genetic evidence of sympatric host races in the seabird tick Ixodes uriae

Due to the close association between parasites and their hosts, many ‘generalist’ parasites have a high potential to become specialized on different host species. We investigated this hypothesis for a common ectoparasite of seabirds, the tick Ixodes uriae that is often found in mixed host sites. We examined patterns of neutral genetic variation between ticks collected from Black‐legged kittiwakes (Rissa tridactyla) and Atlantic puffins (Fratercula arctica) in sympatry. To control for a potential distance effect, values were compared to differences among ticks from the same host in nearby monospecific sites. As predicted, there was higher genetic differentiation between ticks from different sympatric host species than between ticks from nearby allopatric populations of the same host species. Patterns suggesting isolation by distance were found among tick populations of each host group, but no such patterns existed between tick populations of different hosts. Overall, results suggest that host‐related selection pressures have led to the specialization of I. uriae and that host race formation may be an important diversifying mechanism in parasites.     

Pereskiophaga brasiliensis,a natural enemy of the invasive alien cactus Pereskia aculeata,is not suitably host specific for biological control in South Africa

The stem-mining weevil, Pereskiophaga brasiliensis, was a candidate biological control agent for the invasive cactus Pereskia aculeata in South Africa. In host specificity trials, it developed on two indigenous test plant species under choice and no-choice conditions. Pereskiophaga brasiliensis is therefore not suitably host specific for release in South Africa.     

Should the mirid,Eccritotarsus catarinensis (Heteroptera: Miridae), be considered for release against water hyacinth in the United States of America?

Between one and seven biological control agents have been released against water hyacinth (Eichhornia crassipes (Mart.) Solms) in at least 30 countries, with varied success. A mirid, Eccritotarsus catarinensis (Carvalho) (Heteroptera: Miridae), the most recent agent released, is damaging to the plant on the African continent. It could be useful in the USA where water hyacinth remains a problem, but its introduction remains in doubt because during host specificity trials, it developed on Pontederia cordata L. (pickerelweed), indigenous to the USA. However, it did not establish on pickerelweed monocultures during South African field trials, and only light spillover feeding occurred where the two plants coexisted suggesting that the use of P. cordata as a host is a laboratory artefact and it may be suitable for use in the USA, if its thermal physiology allows establishment. We reran models developed for South Africa using CLIMEX to predict whether the mirid will establish where water hyacinth and pickerelweed co-occur, but not where pickerelweed occurs in the absence of water hyacinth. The models suggest that the mirid's distribution will be limited by cold winter temperatures and insufficient thermal accumulation to the southern states of the USA, within the main distribution of water hyacinth. Even though some spillover feeding on pickerelweed might result where the two plants co-occur, the risk of population level effects seems minimal and the risk to more northern pickerelweed negligible. The benefits, including improved habitat for pickerelweed, associated with further suppression of water hyacinth, outweigh the minimal risk of collateral damage to pickerelweed.     

Ancient and modern colonization of North America by hemlock woolly adelgid,Adelges tsugae (Hemiptera: Adelgidae), an invasive insect from East Asia

Hemlock woolly adelgid, Adelges tsugae, is an invasive pest of hemlock trees (Tsuga) in eastern North America. We used 14 microsatellites and mitochondrial COI sequences to assess its worldwide genetic structure and reconstruct its colonization history. The resulting information about its life cycle, biogeography and host specialization could help predict invasion by insect herbivores. We identified eight endemic lineages of hemlock adelgids in central China, western China, Ulleung Island (South Korea), western North America, and two each in Taiwan and Japan, with the Japanese lineages specializing on different Tsuga species. Adelgid life cycles varied at local and continental scales with different sexual, obligately asexual and facultatively asexual lineages. Adelgids in western North America exhibited very high microsatellite heterozygosity, which suggests ancient asexuality. The earliest lineages diverged in Asia during Pleistocene glacial periods, as estimated using approximate Bayesian computation. Colonization of western North America was estimated to have occurred prior to the last glacial period by adelgids directly ancestral to those in southern Japan, perhaps carried by birds. The modern invasion from southern Japan to eastern North America caused an extreme genetic bottleneck with just two closely related clones detected throughout the introduced range. Both colonization events to North America involved host shifts to unrelated hemlock species. These results suggest that genetic diversity, host specialization and host phylogeny are not predictive of adelgid invasion. Monitoring non‐native sentinel host trees and focusing on invasion pathways might be more effective methods of preventing invasion than making predictions using species traits or evolutionary history.