Host compatibility interacts with seed dispersal to determine small-scale distribution of a mistletoe in Xishuangbanna,Southwest China

Aims Mistletoe infection between intra- and interspecific hosts can be restricted by seed dispersal, host–mistletoe compatibility and other factors, yet few studies have linked seed dispersal and seedling establishment together for understanding mistletoe plant distribution and demography together in different anthropogenic disturbance forest types at a local scale. The objectives of this study were to examine how three factors—seed disperser behavior, post-dispersal host compatibility and canopy cover—affect the spatial distribution of a generalist mistletoe Dendrophthoe pentandra (Loranthaceae) in plantation and rainforest within Xishuangbanna, Southwest China.Methods We observed mistletoe D. pentandra infection patterns at the scale of individual trees and sixteen 400-m 2 forest plots in adjacent plantation and rainforest within Xishuangbanna. To elucidate what determines infection patterns at different scales and in different forest types, we observed the behavior of major avian seed dispersers and carried out a seed inoculation experiment to examine how post-dispersal compatibility and light incidence affect the infection of different hosts.Important findings Dendrophthoe pentandra displayed an aggregated distribution and infected 10 species in our study site, with a significantly higher infection prevalence and intensity in the plantation than in the tropical forest. Different seed dispersers provided contrasting initial mistletoe templates: the specialist frugivore Dicaeum concolor (plain flowerpecker) preferred to fly between mistletoes in infected trees in the plantation and likely intensified existing infections. In contrast, the dietary generalist Pycnonotus jocosus (red-whiskered bulbul) was more likely to visit uninfected trees, thereby establishing new infections. Thus, seed dispersal appears to be an important determinant of the mistletoes distribution, with deposition patterns providing an initial distribution template and determining small-scale patterns. However, post-dispersal and abiotic factors revealed that different host compatibilities and levels of light incidence in different habitats affected the survival of D. pentandra seedlings. Hence, our findings suggest that seed dispersal interacts with host compatibility and canopy cover to determine establishment success, survival and the observed distribution patterns.     

Distribution and dispersal of desert mistletoe is scale-dependent, hierarchically nested

Spatial patterns are important to many ecological processes, and scale is a critical component of both patterns and processes. I examined the pattern and scale of the spatial distribution of infection of host plants by the desert mistletoe, Phoradendron californicum, in a landscape that spans several square kilometers. I also studied the relationship between mistletoe infection and seed dispersal. I found elevated seed rain in areas with a high prevalence of mistletoes and I found that a greater proportion of trees receive seeds than are infected, suggesting that mistletoes will be aggregated in space. Using nested analysis of variance and variograms, I found that mistletoe infections were distributed in hierarchical patches. Mistletoes were aggregated within trees and mistletoe prevalence was correlated at scales of <1500 m, and at scales >4000 m. Patterns at the largest scales were correlated with elevation: sites at higher elevations showed reduced mistletoe infection compared to those at lower elevations. I propose that at small scales, mistletoe distributions are primarily the result of aggregation of seed-dispersing birds, and that the elevational effect could reflect the recent colonization of higher elevations by the mistletoes' mesquite hosts or the limits of the mistletoes' physiological tolerance to freezing-induced cavitation.     

Implications of movement patterns of a dietary generalist for mistletoe seed dispersal

Mistletoes are dispersed primarily by frugivorous birds and have highly aggregated distributions at multiple scales. Mistletoe specialist frugivores have been found to intensify infections within infected hosts and stands, and this is considered the most likely mechanism underlying clumped mistletoe distributions at these scales. How these patchy infections first develop and whether seed dispersers also contribute to aggregated mistletoe distributions at landscape and regional scales have not been evaluated. Here we predict the mistletoe seed shadow of a dietary generalist (spiny‐cheeked honeyeater Acanthagenys rufogularis Aves: Meliphagidae), by combining our observations of movements via radio telemetry with previous data on gut passage times to estimate seed dispersal curves for individual birds. There was considerable variation in movements and inferred seed dispersal between individuals, with non‐breeding birds predicted to regularly transport Amyema quandang (Santalales: Loranthaceae) seeds up to 700 m; well beyond the boundaries of an existing mistletoe infection. As the first work to consider explicitly the distance component of mistletoe seed dispersal by dietary generalists, this study poses further questions about the relative seed dispersal roles of dietary generalists and mistletoe specialists. Moreover, our findings highlight considerable intraspecific variation in movement and foraging behaviour, suggesting gender and reproductive status of birds should be considered explicitly when quantifying seed dispersal services.     

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.     

Parasites boost productivity: effects of mistletoe on litterfall dynamics in a temperate Australian forest

The importance of litter in regulating ecosystem processes has long been recognised, with a growing appreciation of the differential contribution of various functional plant groups. Despite the ubiquity of mistletoes in terrestrial ecosystems and their prominence in ecological studies, they are one group that have been overlooked in litter research. This study evaluated the litter contribution from a hemiparasitic mistletoe, Amyema miquelii (Lehm. ex Miq.) Tiegh., in an open eucalypt forest (Eucalyptus blakelyi, E. dwyeri and E. dealbata), at three scales; the forest stand, single trees and individual mistletoes. Litter from mistletoes significantly increased overall litterfall by up to 189%, the amount of mistletoe litter being proportional to the mistletoe biomass in the canopy. The high litter input was due to a much higher rate of mistletoe leaf turnover than that of host trees; the host litterfall and rate of leaf turnover was not significantly affected by mistletoe presence. The additional litter from mistletoes also affected the spatial and temporal distribution of litterfall due to the patchy distribution of mistletoes and their prolonged period of high litterfall. Associated with these changes in litterfall was an increase in ground litter mass and plant productivity, which reflects similar findings with root-parasitic plants. These findings represent novel mechanisms underlying the role of mistletoes as keystone resources and provide further evidence of the importance of parasites in affecting trophic dynamics.     

Ecological assembly rules on arthropod community inhabiting mistletoes

1. The colonisation of a new habitat by a community is led by deterministic and stochastic processes at different spatio-temporal scales. Parasitic plants, such as mistletoe, represent a new habitat within forest canopy that is free to be colonised by many organisms. 2. This study investigates how ecological factors operating at forest and plant scales drive changes in both specialist (mistletoe-dwelling) and tourist (transient visitors) arthropod communities inhabiting European mistletoe, Viscum album subsp. austriacum, in a Mediterranean pine forest. The influence of elevation along a broad elevational gradient was tested by sampling arthropod communities dwelling in mistletoe plants and host pine branches and the effects of mistletoe plant size, distances to other mistletoes, and temporal variation in arthropod assemblages inhabiting mistletoes. 3. The diversity of the specialist community remained constant along the elevational gradient and over the summer period, while the tourist and pine-dwelling arthropod communities showed species turnover. Larger mistletoes were occupied by more species and individuals, whereas more isolated mistletoes presented the same equilibrium point as the more aggregated ones. Thus, mistletoe size is key to the composition of the arthropod community. 4. In conclusion, this study's findings indicate contrasting assembly rules for specialised and tourist arthropod communities associated with mistletoe. The specialist community was highly stable and followed a deterministic trophic sequence of colonisation as the assemblage rule: first, colonisation by the main specialist herbivore, Cacopsylla visci, and, second, by its predator Anthocoris visci. Meanwhile, the tourist community, being a subset of the arthropod assemblage of the pine, acts independent of mistletoe presence.     

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.     

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.     

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.     

Spatial patterns and interspecific associations of dominant tree species in two old-growth karst forests,SW China

Spatial patterns and interspecific associations of plant species in forests are important for revealing how species interact with each other and with the environment, and hence have important implications for optimal forest management and restoration in degraded forest ecosystems. In this paper, the O-ring statistics were used to characterize the spatial patterns and interspecific associations of eight dominant tree species in two 1-ha old-growth karst forest plots in Maolan National Natural Reserve, southwestern China. We found that most of the eight dominant tree species in two forests were continuously regenerating populations. Six species (Platycarya longipes, Acer wangchii, Clausena dunniana, Castanopsis carlesii var. spinulosa, Distylium myricoides, and Rhododendron latoucheae) exhibited significant aggregations at the majority of scales while others (Celtis biondii and Cyclobalanopsis myrsinaefolia) showed a random distribution pattern at most scales. Negative association was a dominant pattern for most species pairs in the two plots, while positive associations were found at most scales for only two species pairs (Platycarya–Clausena and Castanopsis–Rhododendron). Results also indicated that the two main factors of habitat complexity and heterogeneity—the elevation and rock-bareness rate—play important roles in determining spatial distribution patterns and interspecific associations of tree species in karst forests of Maolan. Thus, the observed spatial patterns among the eight tree species are influenced by habitat heterogeneity in the context of karst topographical variations. The partitioning of habitat niches contributes to the promoting species coexistence in species-rich karst forests. The differences of species features in spatial patterns and associations should be paid more attention when planning forest management and developing restoration strategies.     

Spatial patterns at host and forest stand scale and population regulation of the pine processionary moth Thaumetopoea pityocampa

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Parasite-induced parthenogenesis in a freshwater snail: stable,persistent patterns of parasitism

Summary The role of parasites in the evolution of host reproductive modes has gained renewed interest in evolutionary ecology. It was previously argued that obligate parthenogenesis (all-female reproduction) arose in a freshwater snail, Campeloma decisum, as a consequence of severe sperm limitation caused by an unencysted trematode, Leucochloridiomorpha constantiae. In the present study, certain conditions are examined for parasitic castration to account for the maintenance of parthenogenesis: the spatial patterns of the prevalence and intensity of infection on a broad geographical scale and its relationship to host genotype; the recovery from infection after isolation from sources of infection; age-related patterns of infections; and the effects of L. constantiae on snail fecundity.In contrast to the common pattern of the aggregated distribution of parasites within host populations, many snail populations with high prevalence and intensity of infection have non-aggregated parasite distributions. Clonal genotype of the host explained little of the variation in intensity and prevalence of infection by the parasite. Female snails maintained similar prevalence and intensity of infection after isolation, and individuals accumulated parasites throughout their lifespan, both of which suggest there is no effective immune response to infection by L. constantiae. Snail fecundity is not significantly influenced by the intensity of infection. These results suggest that L. constantiae may have represented a strong selective force against males during the initial introduction of this parasite into sexual snail populations because of the persistent nature of infection.     

择伐对思茅松自然种群结构和空间分布格局的影响

思茅松是云南省重要的材脂兼用树种,而择伐是思茅松天然林管理的有效手段,也是获取木材的重要途径。基于对云南省景谷县思茅松天然林的样地调查数据,采用种群径级结构代替年龄结构、Ripley的L函数点格局等方法,对思茅松天然林在不同强度择伐10a后的种群结构和空间分布格局及其不同发育阶段在不同尺度下的相互关系进行了系统研究和分析。结果表明:择伐10a后3种类型的思茅松自然种群的胸径、树高之间呈显著的正相关且可用幂指数模拟预测模型。未择伐的思茅松种群呈衰退型,轻度择伐(12.9%)后种群呈稳定型,中度择伐(29.3%)后的种群属增长型,适当的中度择伐有利于自然种群的更新;未择伐与中度择伐后的思茅松种群分布格局整体呈随机型,但后者较前者更趋向于均匀分布,轻度择伐后的思茅松种群分布格局随着尺度的增加由随机转向聚集分布。择伐对不同径级思茅松天然林种群空间分布格局有重要影响,未择伐的思茅松自然种群随着发育阶段的发展,多数尺度上由随机分布逐渐转变为聚集分布,至大树则又以随机分布为主;轻度择伐和中度择伐的思茅松自然种群则随着发育阶段的发展而由完全的随机分布逐渐转变为聚集分布,中度择伐的聚集分布则更为明显。思茅松不同发育阶段间在所有尺度上基本都呈显著正相关,表明思茅松种群种内竞争弱,有利于种群的维持。     

Identifying the abiotic and biotic drivers behind the elevational distribution shift of a parasitic plant

• Climate change will alter the biotic and abiotic environment and dissipate ecological barriers, reorganising maps of current distribution of parasites and their hosts. In this study, we analyse the population dynamics of the parasitic plant Viscum album subsp. austriacum and explore key biotic (host availability and seed dispersal) as well as abiotic (temperature) factors influencing elevational distribution.
• The study was conducted along an elevational gradient of a Mediterranean mountain, covering the distribution belts of three potential pine hosts: Pinus halepensis (1300–1500 m), P. nigra (1300–1900 m) and P. sylvestris var. nevadensis (1600–2000 m). Along this gradient, we measured multiple variables of mistletoe population (prevalence, abundance and demographic profile) and different factors that might define the current mistletoe distribution (host suitability and availability, temperature and seed dispersal services).
• We found a decline in mistletoe prevalence and abundance with increasing elevation, detecting larger values of both variables at lower elevations of the most suitable host (Pinus nigra). Pinus sylvestris var. nevadensis was a suboptimal but suitable host for the parasite at high elevations. Mistletoe found suitable temperatures and seed dispersal services all along the gradient, being able to recruit at any site.
• With warming temperatures, the presence of suitable vectors for parasite dispersion, and the presence of a sub‐optimal host (P. sylvestris var. nevadensis) at the mountain top, mistletoe currently has a window of opportunity to expand its present geographic distribution to the summits.

     

Directed dispersal of mistletoe (Plicosepalus acaciae) by Yellow-vented Bulbuls (Pycnonotus xanthopygos)

We studied avian dispersal of seeds from the hemiparasitic mistletoe Plicosepalus acaciae (Loranthaceae) to its tree hosts Acacia raddiana and A. tortilis in the Syrian–African Rift (Arava) valley, Israel. The Yellow-vented Bulbul (Pycnonotus xanthopygos) was the sole avian visitor observed feeding on mistletoe fruits. Bulbuls consumed mistletoe fruits whenever they were available, but the fruits only constituted a significant portion of the diet (71% of foraging attempts) when they were most abundant. These birds are potentially good dispersal vectors of P. acaciae because they swallowed the fruit whole and defecated viable seeds that were covered in a viscid pulp, which allowed the seeds to adhere to substrates when voided. In addition, bulbuls spent a large proportion (66–93%) of total observation time perched in Acacia trees, allowing for directed dispersal. Ephemeral river valleys (wadis) with high mistletoe infection were adjacent to those containing no infections, demonstrating that mistletoe dispersal is common within, but not among wadis. This is consistent with the flight behaviour in bulbuls, which do not typically move among wadis. We combined data on bulbul movements between Acacia trees with transit times of mistletoe seeds to create a hypothetical seed shadow as a function of distance from the parent mistletoe plant. Because they are directed dispersers, the movement patterns of bulbuls may explain the current distribution of P. acaciae in the Arava valley.     

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.