Biochemical and morphological evidence for host race evolution in desert mistletoe,Phoradendron californicum (Viscaceae)

Allozymes and morphological characters were used to test whether host race evolution—the genetic divergence of parasitic populations caused by adaptation to different host species—has occurred in desert mistletoe,Phoradendron californicum. Populations ofPhoradendron californicum from two hosts,Acacia greggii andProsopis glandulosa, were surveyed from the Mojave and Colorado deserts. Electrophoretic data indicated genetic differentiation of mistletoes occurring on these hosts. Three of four morphological characters (internode length, main shoot lateral shoot diameter ratio and berry color) also showed significant host-specific differentiation. These data support the hypothesis that host race formation has occurred or is occurring in this parasitic angiosperm.     

Do differences in life‐history traits and the timing of peak mating activity between host‐associated populations of Chilo suppressalis have a genetic basis?

The development of host races, genetically distinct populations of the same species with different hosts, is considered to be the initial stage of ecological speciation. Ecological and biological differences consistent with host race formation have been reported between water‐oat and rice‐associated populations of Chilo suppressalis. In order to confirm whether these differences have a genetic basis, we conducted experiments to determine the extent to which various life‐history traits and the time of peak mating activity of these populations were influenced by the species of host plant larvae were raised on. Individuals from each population were reared for three consecutive generations on either water‐oat fruit pulp or rice seedlings. Descendants of both populations had higher larval survival rates, shorter larval developmental periods, higher pupal weight, and longer adult forewings, when reared on water‐oats than when reared on rice. The time of peak of mating activity differed between the descendants of each population, irrespective of whether they were raised on water‐oats or rice. These results indicate that although some life‐history traits of host‐associated populations of C. suppressalis are influenced by the host plant larvae are raised on, time of peak mating activity is not. Because it is a stable, objective, phenotypic trait, further research on difference in the time of peak mating activity between host‐associated populations of C. suppressalis should be conducted to clarify the mechanism responsible for host race formation in this species.     

Modelling the evolution of common cuckoo host‐races: speciation or genetic swamping?

Co‐evolutionary arms races have provided clear evidence for evolutionary change, especially in host–parasite systems. The evolution of host‐specific races in the common cuckoo (Cuculus canorus), however, is also an example where sexual conflict influences the outcome. Cuckoo females benefit from better adaptation to overcome host defences, whereas cuckoo males face a trade‐off between the benefits of better adaptation to a host and the benefits of multiple mating with females from other host‐races. The outcome of this trade‐off might be genetic differentiation or prevention of it by genetic swamping. We use a simulation model to test which outcome is more likely with three sympatric cuckoo host‐races. We assume a cost for cuckoo chicks that express a host adaptation allele not suited to their foster host species and that cuckoo males that switch to another host‐race experience either a fitness benefit or cost. Over most of the parameter space, cuckoo male host‐race fidelity increases significantly with time, and gene flow between host‐races ceases within a few thousand to a hundred thousand generations. Our results hence support the idea that common cuckoo host‐races might be in the incipient stages of speciation.     

The Relative Contribution of Specialists and Generalists to Mistletoe Dispersal: Insights from a Neotropical Rain Forest

Mistletoes rely on birds for seed dispersal, but the presumed importance of mistletoe‐specialist frugivores has not been critically examined nor compared with generalist frugivores and opportunistic foragers. The contribution of these three groups was compared directly by quantifying bird visitation to fruiting mistletoe plants ( Oryctanthus occidentalis: Loranthaceae) at Barro Colorado Island, Panama, and by comparing these results with proportions calculated from other empirical studies of mistletoe visitation conducted elsewhere. After more than 100 h of timed watches, 23 bird species were recorded visiting eight heavily infected host trees ( Luehea seemannii: Tiliaceae). Eight of these species visited mistletoe, of which five (all tyrannids) consumed mistletoe fruit. Although two mistletoe specialist frugivores ( Tyrannulus elatus and Zimmerius vilissimus) removed most fruit (73%), more than a quarter was consumed by one generalist frugivore ( Mionectes oleagineus) and two opportunists ( Myiozetetes cayanensis and Myiozetetes similis). Post consumption behaviour varied: the specialists flew from mistletoe to mistletoe, the generalist rested in the subcanopy and understory, and the opportunists spent most time hawking insects and resting high in the canopy. Integrating these data with previous work, the dietary specialization, short gut passage rate and strict habitat preferences of mistletoe specialists suggests that their services relate primarily to intensification and contagious dispersal, while species with broader diets are more likely to visit uninfected trees and establish new infections. The presumed importance of mistletoe‐specialist frugivores was not supported and mistletoes are considered to be comparable to many other bird‐dispersed plants, relying on both specialist and generalist frugivores, while opportunists may be disproportionately important in long‐distance dispersal.     

Effectiveness of Mistletoe Seed Dispersal by Tyrant Flycatchers in a Mixed Andean Landscape

The dependence of mistletoes on few dispersers and the directed dispersal they provide is well known, yet no recent work has quantified either the effectiveness of these ‘legitimate’ dispersers, or the extent of redundancy among them. Here, I use the seed dispersal effectiveness (SDE) framework to analyze how birds (Mionectes striaticollis and Zimmerius bolivianus) contribute to mistletoe (Struthanthus acuminatus and Phthirusa retroflexa) infection in traditional mixed plantations within a humid montane forest in Bolivia. I calculated SDE for each bird–mistletoe pair and for the disperser assemblage, by estimating both the quantity and the quality of dispersal. The quantity of dispersal was measured as: (1) disperser abundance; (2) frequency of visits; and (3) number of seeds dispersed per visit, and the quality of dispersal was measured as: (1) germination percentage and speed of germination of seeds regurgitated by birds; and (2) the concordance of deposited seeds and seedling distribution patterns with adult mistletoe distribution at three scales (habitat, host, and microhabitat). Dispersers were not redundant: the more generalist species M. striaticollis dispersed more seeds, but provided lower quality seed dispersal, whereas the mistletoe specialist Z. bolivianus provided low‐quantity and high‐quality seed dispersal. Whereas S. acuminatus benefited more from the SDE of Z. bolivianus, P. retroflexa benefited from the complementary seed dispersal provided by both birds. These results demonstrate how sympatric mistletoes that share the same disperser assemblage may develop different relationships with specific vectors, and describe how the services provided by two different dispersers (one that provides high‐quality and one that provides high‐quantity dispersal) interact to shape spatial patterns of plants.     

A Host Creates an Enemy‐free Space for Mistletoes by Reducing Seed Predation Caused by a Woodboring Beetle: A Hypothesis

Parasitic plants rely on host plants for nutrition. The number of host species varies largely between groups, from single species or genus to hundreds of species. Relative abundance of the host and evolutionary history are the main requisites for parasitic plants to develop specificity to abundant hosts. In the present study, we suggest a novel mechanism by which a hemiparasitic mistletoe can develop local specificity mediated by its host. First, we describe a novel interaction in which a woodboring beetle (Hypothenemus obscurus) preys on mistletoe seeds (Psittacanthus plagiophyllus) attached to tree branches. This beetle preys more frequently on seeds deposited on branches of non‐host species than on branches of its unique local host species (53 percent on average vs. 26 percent respectively). We hypothesize that local host specificity for this mistletoe could be partly mediated by beetle‐host incompatibility, since the host offers a predation‐free space in which mistletoes have better chances to grow. Furthermore, that the exceptional gum exudates produced by this unique host species minimize beetle attacks on branches, thus reducing predation of mistletoe seeds. This novel tritrophic interaction opens an avenue for research on macroscopic host‐specificity mechanisms that occur at the level of the host bark and that have been poorly studied by plant ecologists.     

Mistletoe macroecology: spatial patterns in species diversity and host use across Australia

Mistletoes are parasitic plants, the spatial distributions of which are poorly understood on macroecological scales. Because of their highly unusual life history, investigating mistletoe macroecology may provide new insight into broad‐scale patterns in species distributions. We collated data on the spatial distribution and host use of 65 species of Loranthaceous mistletoes across the continent of Australia, and tested two predictions. First, we predicted mistletoe diversity would be unrelated to productivity (i.e. evapotranspiration and precipitation), as the parasitic lifestyle might relax environmental constraints on their distributions. Second, we predicted that mistletoe host ranges (number of infected host species) would increase in areas with more potential host species. The basis of this prediction is that greater host generality is likely to evolve in regions with greater host diversity because of greater unpredictability in encounter rates with particular host species. Conversely, in regions with fewer potential hosts, randomly dispersing mistletoe propagules are likely to repeatedly encounter particular host species, thus favouring the evolution of host specialization. The results were generally consistent with these predictions. Mistletoe diversity across Australia was weakly associated with environmental conditions, whereas mistletoe host ranges increased significantly with total plant diversity. Macroecological patterns in mistletoes are unusual. In contrast to non‐parasitic plants, mistletoe diversity is poorly correlated with productivity. Host ranges varied predictably across Australia, providing the first quantitative support for the hypothesis that mistletoes in diverse regions tend to be host generalists, whereas mistletoes in depauperate regions tend to be host specialists. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 459–468.     

Mistletoes Play Different Roles in a Modular Host–Parasite Network

Antagonistic interactions between host plants and mistletoes often form complex networks of interacting species. Adequate characterization of network organization requires a combination of qualitative and quantitative data. Therefore, we assessed the distribution of interactions between mistletoes and hosts in the Brazilian Pantanal and characterized the network structure in relation to nestedness and modularity. Interactions were highly asymmetric, with mistletoes presenting low host specificity (i.e., weak dependence) and with hosts being highly susceptible to mistletoe‐specific infections. We found a non‐nested and modular pattern of interactions, wherein each mistletoe species interacted with a particular set of host species. Psittacanthus spp. infected more species and individuals and also caused a high number of infections per individual, whereas the other mistletoes showed a more specialized pattern of infection. For this reason, Psittacanthus spp. were regarded as module hubs while the other mistletoe species showed a peripheral role. We hypothesize that this pattern is primarily the result of different seed dispersal systems. Although all mistletoe species in our study are bird dispersed, the frugivorous assemblage of Psittacanthus spp. is composed of a larger suite of birds, whereas Phoradendron are mainly dispersed by Euphonia species. The larger assemblage of bird species dispersing Psittacanthus seeds may also increase the number of hosts colonized and, consequently, its dominance in the study area. Nevertheless, other restrictions on the interactions among species, such as the differential capacity of mistletoe infections, defense strategies of hosts and habitat types, can also generate or enhance the observed pattern. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

Host-mediated effects on the reproductive phenological asynchrony of a generalist mistletoe in China

一种泛性桑寄生植物繁殖物候异步性的寄主介导效应 寄主介导效应被认为会导致半寄生性的桑寄生植物的繁殖物候异步性，并由此为与桑寄生植物互惠共生的传粉者和种子散布者提供更长时间的食物资源供应，但目前关于此方面的研究还缺乏相关的实证数据。本研究以广泛分布于中国西南西双版纳地区的一种泛性桑寄生科植物五蕊寄生(Dendrophthoe pentandra)为材料，每周监测其开花、结果物候，检测了其开花和结果物候是否呈季节性格局，量化了五蕊寄生繁殖物候的异步程度，并检测了影响该植物始花期早晚的因素。最后，本研究还检验了五蕊寄生繁殖物候的异步性随寄主种类数量变化的效应。研究结果表明：(i)在连续两年的物候观测中五蕊寄 生的花期和果期都呈单峰分布格局；(ii)始花期显著受到植物大小和光照强度的影响，即冠幅越大和受光程度越高的植物个体有更早的始花期和更长的花期和果期；(iii)不同的寄主种类对五蕊寄生的繁殖物候有显著的影响，但与假设相反的是，随着寄主种类数量的增加，五蕊寄生繁殖物候的异步性没有显著提高。这项研究表明，在解释泛性桑寄生植物的繁殖物候异步性及寄主种类的数量对其影响的生态学意义还需进行更深入的探究。     

HOST-INDUCED ASSORTATIVE MATING IN HOST RACES OF THE LARCH BUDMOTH

Abstract The likelihood of sympatric speciation is enhanced when assortative mating is a by‐product of adaptation to different habitats. Pleiotropy of this kind is recognized as important in parasites that use their hosts as a long‐range cue for finding mates, but is generally assumed to have limited applicability for most other organisms. In the larch budmoth, Zeiraphera diniana (Lepidoptera: Tortricidae), sympatric host races feed on larch or pine. Zeiraphera diniana females attract males (call) by releasing host‐independent long‐range pheromones. Pheromone composition differs strongly between host races, but we show in an experimental field study that cross‐attraction can occur at a rate of 0.03–0.38. Cross‐attraction to larch females increases when they call from neighborhoods (8‐m radius) rich in pine or from pine trees. Cross‐attraction to pine females similarly increases when calling from neighborhoods rich in larch, but there is no significant effect of calling substrate. Males, as well as females, of this species preferentially alight on their own host, and in neighborhoods where their own host is common. This effect of tree species and host neighborhood on assortative mating is therefore due, at least in part, to the numbers of males of each host race present within approximately 200 m² surrounding the female. This proximity effect is enhanced by the clumped distributions of the hosts themselves. Host chemistry might also affect pheromone production and/or response directly, but we have evidence neither for nor against this. This work provides empirical evidence that host adaptation has a pleiotropic effect on assortative mating in a species with host‐independent long‐range mating signals. Sympatric speciation via pleiotropy between ecological traits and assortative mating may thus be more common than generally supposed: Clumped resource distributions and habitat choice by adults are widespread.     

Uncorrelated mistletoe infection patterns and mating success with local host specialization in <Emphasis Type="Italic">Psittacanthus calyculatus</Emphasis> (Loranthaceae)

Mistletoe infection between conspecific and interspecific hosts can be restricted by seed dispersal, host-mistletoe compatibility and abiotic factors, yet no studies have linked mistletoe infection patterns and pollination together for understanding mistletoe distribution at a local scale. Psittacanthus calyculatus (Loranthaceae) is a hemiparasitic plant with a broad host range across its geographic distribution. The potential for local host adaptation has been shown using cross-inoculation experiments, in which plants of mistletoe seeds collected from a given host are more likely to survive when they are inoculated on conspecific host trees compared with those inoculated on other host provenances. Here we evaluate host adaptation by describing the local patterns of infection (prevalence and intensity) of P. calyculatus mistletoes on three native host tree species (Alnus acuminata, Quercus crassipes, Salix bonplandiana) and one introduced species (Populus alba) and carried out cross-pollination experiments to examine how pollination affects infection patterns of different host species. Mistletoe infection prevalence (proportion of infection) and infection intensity (mean number of mistletoes per tree) were in general disproportional with respect to the availability of native host tree species but higher to that of non-native host tree species. Cross-pollination experiments showed higher mating success on the native host tree species, suggesting higher local adaptation to specially Q. crassipes. The observed spatial distribution of host tree species and mistletoe infection along with the non-random mating could contribute to local genetic structuring of mistletoe populations.     

Carbon and nitrogen parasitism by a xylem‐tapping mistletoe (Tapinanthus oleifolius) along the Kalahari Transect: a stable isotope study

The present study explores the xylem‐tapping parasitism by mistletoe (Tapinanthus oleifolius) on native tree species along the Kalahari Transect (KT) using the stable isotopes of carbon and nitrogen. Mistletoe‐host pairs were collected at three geographical locations along the KT rainfall gradient in the 2005 and 2006 wet seasons. Foliar total carbon, total nitrogen and their stable isotope compositions (δ¹³C and δ¹⁵N) were measured. Heterotrophy (H) was calculated using foliar δ¹³C values of mistletoes and their hosts as an indicator of proportion of carbon in the mistletoes derived from host photosynthate. Based on the mistletoe H‐value and relationship between the mistletoe foliar δ¹⁵N and their host foliar δ¹⁵N, the results showed that mistletoes along the KT derived both nitrogen and carbon from their hosts. Mistletoes may regulate water use in relation to nitrogen supply. The proportion of carbon in the mistletoes derived from host photosynthate was between 35% and 78%, and the degree of heterotrophy was species‐specific with only limited annual variation. The study emphasizes the importance of incorporating parasitic associations in future studies on studying carbon, water and nutrient cycling along the Kalahari.     

Host‐associated pre‐mating reproductive isolation between host races of Acrocercops transecta: mating site preferences and effect of host presence on mating

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The contribution of mistletoes to nutrient returns: Evidence for a critical role in nutrient cycling

Both nutrient cycling and nutrient relationships between mistletoe and host have been widely studied; yet it is unclear whether high nutrient concentrations commonly found in mistletoes affect rates of nutrient cycling. To address this question, we assessed 13 elements in the leaf litter of a temperate eucalypt forest in southern New South Wales, comparing concentrations from trees (Eucalyptus blakelyi, E. dwyeri, and E. dealbata) with and without the hemiparasitic mistletoe Amyema miquelii. Results were in accord with previous research on fresh leaves showing that concentrations of many elements were higher in the mistletoe than the host. This was not the case for all elements; most notably for N, where concentrations were significantly lower in the mistletoe. However, the return of all elements increased with mistletoe infection because of the combined effect of enrichment in mistletoe tissues and high rates of mistletoe litterfall. Annual returns of N and P in leaf litter increased by a factor of 1.65 and 3 respectively, with the greatest increase being for K by a factor of 43 in spring. These increased element returns were not significantly influenced by any changes in host leaf litter quality, as mistletoe infection was not found to affect host element concentrations. Mistletoe infection also altered the spatial and temporal distribution of element returns because of the patchy occurrence of mistletoes and extended period of mistletoe litterfall compared with the host. These findings provide a mechanistic explanation for the role of mistletoes as a keystone resource and, together with comparable results from root‐parasitic plants in boreal tundra and cool‐temperate grasslands, suggest that enhancing nutrient return rates may be a generalized property of parasitic plants.     

Macroscale analysis of mistletoe host ranges in the Andean‐Patagonian forest

• The number of host species infected by a mistletoe (host range) is critical in that it influences prevalence, virulence and overall distribution of the parasite; however, macroecological analyses of this life history feature are lacking for many regions.
• The Andean‐Patagonian forest, found along the southern Andes from 35 °S to Tierra del Fuego at 55 °S, contains 12 mistletoe species in three families (Loranthaceae, Misodendraceae and Santalaceae). By tabulating herbarium records, the host ranges and geographical distributions of these mistletoes were explored.
• Our results show that these parasites occur on 43 plant species in 24 families but with varying degrees of specificity. All Misodendrum species and Desmaria mutabilis (Loranthaceae) are specialists that use Nothofagus as their primary hosts. Tristerix and Notanthera (Loranthaceae) and Antidaphne and Lepidoceras (Santalaceae) are generalists parasitizing more than six host species from several genera and families. Although many of the mistletoe species are sympatric, there is low overlap in host use.
• Our data show that in the southern South American bioregion, generalist mistletoes have smaller geographic ranges than specialists. This contrast with a previous hypothesis that predicted mistletoes with large geographic ranges would also have large host ranges, and conversely, less diverse regions would have more specialised mistletoes.

     

Consequences of disperser behaviour for seedling establishment of a mistletoe species

The dispersal process in plants links adults and their offspring. For frugivore‐dispersed plants, animal behaviour can have a strong effect on plant fitness. Many mistletoes are totally dependent on animals that deposit seeds on suitable hosts and particular branch diameters. We characterised the seed dispersal and seedling establishment of the mistletoe Tristerix corymbosus, which at our study site, is exclusively dispersed by the marsupial Dromiciops gliroides. Mistletoes’ fruits have a viscous pulp that remains in the seed even after dispersal. This substance adheres the seed to the host branch. We estimated host branch availability in the forest and seed deposition (faeces) by the marsupial in the study area. Specifically, the branch suitability factors we assessed were host identity, branch status (alive or dead), branch diameter, height, and canopy cover. Lodged faeces were individually marked and the number of seed deposited within these droppings was counted, and we recorded the number of seedlings with true leaves that had established after 1 year to estimate the probability of seedling establishment. Branch diameter and canopy cover had a significant positive effect on seed deposition probability. Seedling establishment probability decreased with the number of seeds deposited per faeces and with canopy cover. In general, the marsupial deposited mistletoe seeds in microsites that increase the chance of seedling establishment. Thus, the movement behaviour of the marsupial has a positive effect on the regeneration process of this mistletoe species.     

Host spatial structure and disperser activity determine mistletoe infection patterns

What processes and factors are responsible for species distribution are long-standing questions in ecology and a key element for conservation and management. Mistletoes provide the opportunity to study a forest species whose occurrence is expected to be constrained by multiple factors as a consequence of their life form. We studied the mistletoe Tristerix corymbosus (Loranthaceae) on its most common hosts species in northwest Patagonia. The seeds of this mistletoe are almost exclusively dispersed by the small arboreal and endemic marsupial Dromiciops gliroides (Microbiotheridae). We assessed the underlying causes of plant spatial patterns through point pattern analysis and we used different variables that characterize the neighborhood around each host to analyze the relative effect of host availability, potential for disperser movement and canopy light conditions. We found that potential hosts were strongly aggregated and that the three most common host species were distributed independent of each other. Considering all host species together, infected and non-infected host were individually aggregated but segregated from each other. The aggregated pattern of infected hosts could be explained in part by the template of potential hosts distribution, but was subsequently modulated by the activity of the mistletoe disperser. Potential for disperser movement, the proximity to reproductive mistletoes and habitat complexity, increased mistletoe infection probability. However, neighboring host availability decreased mistletoe infection probability, and tree DBH (used as surrogate for light conditions) had no detectable effect. Our results suggested that the distribution of mistletoe infection was determined by the structure of potential host populations and by the marsupial disperser activity. Compared to bird dispersed mistletoes, the scale of the infection was smaller and the proximity to reproductive mistletoes and habitat complexity were important for seed arrival and infection. The interplay between landscape structure and disperser activity determine the spatial structure of mistletoe future generations.     

Variation in mistletoe seed deposition: effects of intra- and interspecific host characteristics

We investigated differences in host infection by a desert mistletoe, Phoradendron californicum, and examined one of the processes that contributes to these differences: variation in seed deposition among host individuals and species. In the Sonoran Desert, P. californicum parasitizes the sympatric leguminous trees Olneya tesota, Cercidium microphyllum, Prosopis velutina, Acacia constricta, and Acacia greggii. We hypothesized that seed deposition depends on host height and crown architecture. At a site in Arizona, frequency of infection did not reflect host relative abundance. Olneya tesota was parasitized at a higher frequency than expected from its abundance and maintained the highest mistletoe loads per individual host. In contrast, P. velutina was infected less frequently than expected. Infection frequency increased with host tree height for all hosts. Mistletoe seed deposition by avian dispersers differed among host species and was disproportionately high in O. tesota and P. velutina. Seed deposition was higher in infected than in non‐infected host trees, and increased with tree height in O. tesota but not in C. microphyllum. We suspect that increased seed deposition with height in O. tesota may be due to the preference of seed‐dispersing birds for higher perches. Some host tree species, such as C. microphyllum and A. constricta, probably received fewer mistletoe seeds because birds avoid hosts with dense and spiny crowns. Mistletoe populations are plant metapopulations in which host trees are patches and the frequency of infection in each host species/patch type is the result of interspecific differences in the balance between mistletoe colonization and extinction. From this perspective, our study of host use and seed dispersal is a metapopulation study of patch occupancy and propagule distribution among available patch types. Our seed‐dispersal study demonstrates that the mechanisms that create pattern in patchy plant populations can be investigated in mistletoe systems.     

西双版纳地区鞘花的繁殖物候特征及影响因素

桑寄生植物作为关键种和关键性食物资源在生态系统中扮演着重要角色,其鞘花的繁殖物候特征不仅会影响自身的繁殖适合度而且还会影响依赖于其获取食物资源的动物。为了解鞘花的繁殖物候特征及其影响因素,探究其与寄主植物和种子散布者之间的相互作用关系。该研究以西双版纳地区分布的鞘花为对象,通过定期观测鞘花和其寄主植物木荷的繁殖物候,测量它们的生物学特性和温湿度等环境因子,并分析鞘花在个体水平和种群水平上的繁殖物候特征以及寄主植物和温湿度对其繁殖物候的影响。结果表明:(1)鞘花的开花物候属于集中大量开花模式,整个种群的花期和果期的持续时间分别约为20 d和72 d,花期和果期的同步性指数都较高,6月中旬鞘花果实被取食的数量和速率最大,之后逐渐降低。(2)鞘花的始花期与木荷的始花期相关性较高,花期和果期与木荷的繁殖物候基本重叠。(3)每月开花和果熟的个体数量与同期和前一个月的平均温度和相对湿度的相关关系均不显著。总之,桑寄生植物的繁殖物候特征可能受到较多因素的影响,若想全面了解半寄生植物的繁殖物候特征,就要综合考虑生物和非生物等多种因子的共同作用。     

Tracing the colonization and diversification of the worldwide seabird ectoparasite Ixodes uriae

Historical patterns of dispersal and population isolation are key components shaping contemporary genetic diversity across landscapes and require explicit consideration when examining the relative role of different factors in driving the evolution of host specificity in parasitic organisms. In this study, we investigate the worldwide colonization history of a common ectoparasite of seabirds, the tick Ixodes uriae. This tick has a circumpolar distribution across both hemispheres but has repeatedly formed host‐specific races within different regions. By combining mitochondrial and nuclear data, we infer how this species spread to its present‐day distribution and how the colonization process may have affected the geographic and host‐associated structure of this tick within regions. We demonstrate that I. uriae is highly structured at a global scale and isolates into four genetic groups that correspond to well‐defined geographical regions. Molecular dating suggests that the diversification of I. uriae began in the early Miocene (22 Myr) and that this tick colonized most of the southern hemisphere before moving into northern latitudes via two independent routes. However, no relationship between the degree of host race divergence and colonization history was evident, supporting previous hypotheses that host specialization evolves relatively rapidly in this parasite, but does not typically lead to speciation. We discuss the possible historical and contemporary mechanisms of large‐scale dispersal for this ectoparasite and how its biological characteristics may condition current patterns of genetic diversity. More generally, our results illustrate how combining broad‐scale sampling and modern molecular tools can help disentangle complex patterns of diversification in widespread parasites.