A phylogenetic analysis of the British flora sheds light on the evolutionary and ecological factors driving plant invasions

Darwin's naturalization hypothesis predicts that invasive species should perform better in their novel range in the absence of close relatives in the native flora due to reduced competition. Evidence from recent taxonomic and phylogenetic‐based studies, however, is equivocal. We test Darwin's naturalization hypothesis at two different spatial scales using a fossil‐dated molecular phylogenetic tree of the British native and alien flora (ca. 1600 species) and extensive, fine‐scale survey data from the 1998 Countryside Survey. At both landscape and local scales, invasive species were neither significantly more nor less related to the native flora than their non‐invasive alien counterparts. Species invasiveness was instead correlated with higher nitrogen and moisture preference, but not other life history traits such as life‐form and height. We argue that invasive species spread in Britain is hence more likely determined by changes in land use and other anthropogenic factors, rather than evolutionary history. Synthesis. The transition from non‐invasive to invasive is not related to phylogenetic distinctiveness to the native community, but instead to their environmental preferences. Therefore, combating biological invasions in the Britain and other industrialized countries need entirely different strategies than in more natural environments.     

The effects of phylogenetic relatedness on invasion success and impact: deconstructing Darwin's naturalisation conundrum

Darwin's naturalisation conundrum describes the paradox that the relatedness of exotic species to native residents could either promote or hinder their success through opposing mechanisms: niche pre‐adaptation or competitive interactions. Previous studies focusing on single snapshots of invasion patterns have provided support to both sides of the conundrum. Here, by examining invasion dynamics of 480 plots over 40 years, we show that exotic species more closely related to native species were more likely to enter, establish and dominate the resident communities, and that native residents more closely related to these successful exotics were more likely to go locally extinct. Therefore, non‐random displacement of natives during invasion could weaken or even reverse the negative effects of exotic–native phylogenetic distances on invasion success. The scenario that exotics more closely related to native residents are more successful, but tend to eliminate their closely related natives, may help to reconcile the 150‐year‐old conundrum.     

基于功能性状多维性理解外来物种入侵的达尔文归化难题

达尔文归化难题是进行外来物种入侵预测的重要理论依据,然而,达尔文归化假说和预适应假说却预测了2种截然不同的结果。事实上,达尔文归化难题争论的焦点是物种间的差异性还是相似性促进了外来物种的成功入侵,究其原因可能是忽略了功能性状的多维性。所谓功能性状的多维性,就是不同的功能性状可能代表着不同的生态功能轴,外来物种的入侵是多个维度上不同生态过程的综合结果。本研究以现代物种共存理论为基础,构建了一个基于环境过滤和适合度差异2个维度的入侵预测模型框架,不同维度对应着不同的功能性状以及不同的种间相似性关系。该预测模型表明,在环境过滤维度上与本地物种性状趋同,同时,在适合度维度上与本地物种性状趋异的外来物种是潜在的入侵物种,而其危害程度主要取决于本地群落的构建过程。该模型框架可为外来物种入侵预警提供理论依据,也可为生物多样性保护、外来物种的防治与管理等提供实践指导。     

It takes one to know one: Similarity to resident alien species increases establishment success of new invaders

Aim

Darwin's naturalization hypothesis states that dissimilarity to native species may benefit alien species establishment due to empty niches and reduced competition. We here add a new dimension to large‐scale tests of community invasibility, investigating the role that previously established alien species play in facilitating or hindering new invasions in plant communities.

Location

Permanent grasslands across France (including mainland and Corsica), as a receding ecosystem of great conservation importance.

Methods

Focusing on 121 alien plant species occurring in 7,215 vegetation plots, we quantified biotic similarity between new invaders and resident alien species (i.e., alien species with longer residence times) based on phylogenetic and trait distances. Additionally, we calculated distances to native species for each alien species and plot. Using multispecies distribution models, we analysed the influence of these biotic similarity measures and additional covariates on establishment success (presence/absence) of new invaders.

Results

We found that biotic similarity to resident alien species consistently increased establishment success of more recently introduced species. Phylogenetic relatedness to previous invaders had an equally strong positive effect as relatedness to native species. Conversely, trait similarity to natives hindered alien establishment as predicted by Darwin's naturalization hypothesis. These results highlight that various mechanisms may act simultaneously to determine alien establishment success.

Main conclusions

Our results suggest that, with greater similarity among alien species, invasion success increases. Such a pattern may arise either due to actual facilitation among invaders or as a result of weaker competitive interactions among invaders than between native and alien species, leading to an indirect facilitative effect. Alternatively, recent environmental changes (e.g., eutrophication, climate change) may have added new environmental filters. Determining how initial invasions might pave the road for subsequent invasions is crucial for effective multispecies management decisions and contributes a new aspect to our understanding of community assembly.

     

基于功能性状的外来植物入侵预测模型框架构建

预测外来植物的潜在入侵性已成为生物多样性保护研究的重要内容,外来植物与乡土物种间的亲缘关系是预测外来植物能否成功入侵的一个重要途径。然而,达尔文归化难题却预测了两种截然不同的结果(即达尔文归化假说和预适应假说)。该研究解析了达尔文归化难题的内涵,提出了基于功能性状的外来植物与乡土群落间的相似性关系应该是进行外来植物入侵预测的重要切入点,而功能性状的种间分化与种内变异可能是外来植物成功入侵的两种不同生态策略。在此基础上,该研究还通过物种功能性状的多维超体积构建了外来植物与乡土群落间的相似性,提出了基于这种相似性的外来植物入侵预测的研究框架和基本流程。该模型框架的建立有助于理解外来植物的入侵机制,对外来植物的潜在入侵性预测提供了理论依据。当然,要实现外来植物能否成功入侵的准确预测,不仅依赖于功能性状的选择,还要考虑入侵的生境依赖性、空间尺度的重要性以及乡土群落的可入侵性等,未来的研究重点是通过控制实验对该模型进行验证和进一步完善。     

Scale dependence in the phylogenetic relatedness of alien and native taxa

达尔文的归化假说提出,由于生态位的不同,成功建群的外来物种与本地物种的关系不太密切。先前的研究对这一假设有支持也有反对,其中一个原因是外来物种和本地物种在大的空间尺度上有系统发育聚类的倾向,而在细微尺度上存在过度分散的倾向。然而,对于外来物种的系统发育关系如何改变其入侵群落的系统发育结构,以及在何种空间尺度上可能表现出这种影响,人们知之甚少。在本研究中,我们调查被入侵的森林下层植物群落在系统发育上是聚集的还是或过度分散的,亲缘关系如何随空间尺度变化,以及外来物种如何影响下层群落的系统发育模式。在澳大利亚东南部干旱森林的下层群落进行了5个空间尺度(1, 20, 500, 1500和4500 m²)的实地调查。使用两个指标的标准化效应量[(i)平均成对距离和(ii)平均最近分类单元距离]来量化群落与其外来和本地亚群落之间的系统发育关系,并研究系统发育模 式如何随空间尺度变化。研究结果表明,外来物种之间的亲缘关系非常密切,而且这种亲缘关系会随着尺度的增加而增加。在中等空间尺 度下(20–500 m²), 整个群落呈随机分布趋势,而本地物种高度分散,外来亚群落高度聚集。这说明亲缘关系密切的外来物种入侵使群落系统发育结构由过度分散向随机分布转变。外来物种和本地物种在空间尺度上是远亲,这支持了达尔文的归化假说,但只是在系统发育距离被量化为平均最近分类单元距离时成立。外来物种和本地物种的系统发育差异随着空间尺度的增加而增加,这与预期的模式相反。我们的研究结果表明,外来物种强大的系统发育聚类是由人类干预的引入驱动的,牵涉能够成功建群和传播的密切相关的类群。系统发育相关性的尺度依赖模式可能是由火灾和散布等随机过程引起的,这表明竞争和生境过滤并不是分别在小和大尺度上控制系统发育关系的唯一因素。区分不同进化深度的指标很重要,因为不同的指标可以显示不同的尺度依赖模式。     

Congener diversity,topographic heterogeneity and human‐assisted dispersal predict spread rates of alien herpetofauna at a global scale

Understanding the factors that determine rates of range expansion is not only crucial for developing risk assessment schemes and management strategies for invasive species, but also provides important insight into the ability of species to disperse in response to climate change. However, there is little knowledge on why some invasions spread faster than others at large spatiotemporal scales. Here, we examine the effects of human activities, species traits and characteristics of the invaded range on spread rates using a global sample of alien reptile and amphibian introductions. We show that spread rates vary remarkably among invaded locations within a species, and differ across biogeographical realms. Spread rates are positively related to the richness of native congeneric species and human‐assisted dispersal in the invaded range but are negatively correlated with topographic heterogeneity. Our findings highlight the importance of environmental characteristics and human‐assisted dispersal in developing robust frameworks for predicting species' range shifts.     

Is body size a good indicator of fecundity in the genus Thaumetopoea? A story told by two intrageneric Mediterranean forest defoliators

1. Body size correlates with several factors such as reproductive fitness, environmental changes, the quality and quantity of food during critical development stages, and the feeding season. For the Palearctic moths of the genus Thaumetopoea (Lepidoptera; Notodontidae), the larval development is crucial and differs between species according to their feeding season; larvae of the pine processionary moth Thaumetopoea pityocampa (Denis & Schiffermüller 1775) feed during winter while larvae of its congeneric cedar processionary moth Thaumetopoea bonjeani (Powell 1922) develop during summer in North Africa.
2. This discrepancy in lifecycles leads to different reproductive traits such as egg batch length, number of eggs per batch, eggs protection mechanisms and female body size.
3. According to Darwin's fecundity advantage hypothesis (1871), female body size should have a positive influence on reproductive fitness, since larger females supposedly have higher fecundity. The universal allometric scaling phenomenon rule proposed by Rensch (1950) predicts that the degree of sexual size dimorphism tends to decrease with the increase of female body size.
4. Here, two morphometrical parameters that is, body size and scale size, estimated from body measurements of individuals of both species, feeding on the same host Atlantic cedar Cedrus atlantica (Manetti & Carrière 1855) (Pinales; Pinaceae) in Algeria were proposed. The aim was to find out traits that might rule the competition for food and space, in particularly fecundity and body size.
5. Results of the present study highlight a female-biased sexual size dimorphism in both species. The positive correlation between female body size and fecundity shown in this study weakly supports Darwin's hypothesis. Finally, the intrageneric test performed leads to conclude that Rensch's rule does not hold in the considered species.

     

Conservation status of landbirds on Floreana: the smallest inhabited Galápagos Island

On Floreana, the smallest inhabited island in the Galápagos, populations of several species of birds have either been extirpated or, based on anecdotal evidence and small‐scale surveys, are declining. Our objective, therefore, was to conduct a comprehensive survey of landbirds encompassing the entire island during three breeding seasons (2014–2016). We conducted surveys at 59 points in 2014, 257 in 2015, and 295 in 2016. Each survey point was sampled once. We detected 12 species during our surveys. Galápagos Flycatchers, Yellow Warblers, Small and Medium ground‐finches, and Small Tree‐Finches were widely distributed over the entire island. Common Cactus‐Finches and Medium Tree‐Finches had more restricted distributions in the lower or higher parts of the island. Few Dark‐billed Cuckoos (Coccyzus melacoryphus), Paint‐billed Crakes (Neocrex erythrops), Galápagos Doves (Zenaida galapagoensis), and Galápagos Short‐eared Owls (Asio flammeus galapagoensis) were recorded. Small Ground‐Finches and Small Tree‐Finches were found at densities comparable to those on other Galápagos Islands, whereas densities of Galápagos Flycatchers and Yellow Warblers were higher on Floreana than on other islands. Endemic Medium Tree‐Finches were confined to an area of 24 km², mainly in the highlands, but were still widespread and common in their restricted habitat, with the number of territories estimated to be between 3900 and 4700. Of 22 originally occurring landbirds on Floreana, no fewer than 10 species have either been extirpated or are likely to have been extirpated since the arrival of the first human inhabitants. The combined effects of introduced mammals, large‐scale habitat destruction, and direct human persecution were responsible for the extirpation of six species during the 19th century. Three additional species have been extirpated since 1960, likely due to the introduction of the parasitic fly Philornis downsi, and this fly remains a major threat for the remaining bird species. Developing strategies for reducing the impact of these flies on the birds of the Galapagos Islands must be a high priority. In addition, habitat management and restoration, including the control of invasive plants and promotion of native tree species, will be critically important in conserving landbird populations on Floreana.     

The importance of phylogenetic scale in tests of Bergmann's and Rapoport's rules: lessons from a clade of South American lizards

We tested for the occurrence of Bergmann's rule, the pattern of increasing body size with latitude, and Rapoport's rule, the positive relationship between geographical range size and latitude, in 34 lineages of Liolaemus lizards that occupy arid regions of the Andean foothills. We tested the climatic-variability hypothesis (CVH) by examining the relationship between thermal tolerance breadth and distribution. Each of these analyses was performed varying the level of phylogenetic inclusiveness. Bergmann's rule and the CVH were supported, but Rapoport's rule was not. More variance in the data for Bergmann's rule and the CVH was explained using species belonging to the L. boulengeri series rather than all species, and inclusion of multiple outgroups tended to obscure these macroecological patterns. Evidence for Bergmann's rule and the predicted patterns from the CVH remained after application of phylogenetic comparative methods, indicating a greater role of ecological processes rather than phylogeny in shaping the current species distributions of these lizards.     

A practical approach to the study of spatial structure in simple cases of heterogeneous vegetation

Abstract. Spatial heterogeneity is a characteristic of most natural ecosystems which is difficult to handle analytically, particularly in the absence of knowledge about the exogenous factors responsible for this heterogeneity. While classical methods for analysis of spatial point patterns usually require the hypothesis of homogeneity, we present a practical approach for partitioning heterogeneous vegetation plots into homogeneous subplots in simple cases of heterogeneity without drastically reducing the data. It is based on the detection of endogenous variations of the pattern using local density and second‐order local neighbour density functions that allow delineation of irregularly shaped subplots that could be considered as internally homogeneous. Spatial statistics, such as Ripley's K‐function adapted to analyse plots of irregular shape, can then be computed for each of the homogeneous subplots. Two applications to forest ecological field data demonstrate that the method, addressed to ecologists, can avoid misinterpretations of the spatial structure of heterogeneous vegetation stands.     

The potential for birds to disperse the seeds of Acacia cyclops,an invasive alien plant in South Africa

Rooikrans Acacia cyclops is an aggressive invasive tree that threatens natural resources in South Africa. The seeds of A. cyclops have a prominent aril which attracts birds that ingest the seeds and disperse them endozoochorously. Two biological control agents, a Seed Weevil Melanterius servulus and a Flower‐galling Midge Dasineura dielsi, were released on A. cyclops in 1991 and 2002, respectively. Together these agents have substantially reduced seed production and generally far lower numbers of seeds are now available to birds. A consequence of this transition from historically bounteous quantities of seeds to scanty seed availability is that birds may no longer associate with the trees and seed dispersal may be disproportionately reduced. To assess whether this has happened, seed attrition was measured by comparing the amount of seeds that disappeared from two groups of branches, one available to birds and the other enclosed in bird netting. Other types of granivores (mainly field mice) were excluded from both groups of branches with a plastic funnel placed around the stems. Mature seeds were also harvested and fed to caged bird species to determine gut retention times and germination rates of ingested seeds. Attrition rates of seeds showed that birds continue to remove seeds but that only a proportion of the crop is taken. Only two frugivorous species (Knysna Turaco Tauraco corythaix and Red‐winged Starling Onychognathus morio) and two granivorous species (Red‐eyed Dove Streptopelia semitorquata and Laughing Dove Streptopelia sengalensis) ingested A. cyclops seeds during feeding trials. Ingestion by birds enhanced seed germination except for those ingested by Laughing Doves. There were no apparent effects of length of gut passage time and avian body size on seed germination rates. Despite the diminished seed resource due to biological control agents, birds continue to disperse A. cyclops seeds.     

Influence of environmental conditions on sex allocation in the black rhinoceros population of Mkhuze Game Reserve,South Africa

According to the Trivers–Willard and local resource competition (LRC) hypotheses, for species where reproductive success is more variable in one sex, natural selection may lead to a bias in sex allocation of a female’s offspring according to her body condition. The extrinsic modification hypothesis (EMH) suggests that offspring sex can also be influenced by environmental conditions experienced by mothers. We investigated the influence of rainfall, El Niño‐Southern Oscillation (ENSO), population size and burning, in the year before conception and during pregnancy, on sex allocation in the black rhinoceros population of Mkhuze Game Reserve, South Africa, during 1970–2007. Females were more likely to have a male calf as rainfall during pregnancy increased, supporting the Trivers–Willard hypothesis. Also, the probability of having a male calf increased with population size, supporting the LRC hypothesis. Calf sex allocation was not influenced by ENSO. In conclusion, local environmental conditions may influence sex allocation in black rhinoceros, thereby supporting the EMH. Burning and population size may influence sex allocation in black rhinoceros, and yet can be manipulated by managers. Thus, this knowledge can be applied to improve population structure assessments and management regimes, especially in enclosed reserves, which is essential to maintain endangered species’ productivity.     

Species-specific impact of introduced largemouth bass Micropterus salmoides in the Groot Marico Freshwater Ecosystem Priority Area,South Africa

Largemouth bass Micropterus salmoides are among the world's 100 worst invaders and negatively affect aquatic biodiversity in many regions worldwide. In South Africa there is a paucity of empirical studies describing their impacts. The impact of M. salmoides on the fish community in the Groot Marico River catchment, an otherwise near-pristine river ecosystem and a freshwater ecosystem priority area, was assessed from surveys conducted in 2012. Fish presence and abundance were enumerated using multiple survey techniques, and their association with key habitat variables and the presence or absence of M. salmoides were assessed. A total of 14 native fish species were recorded, besides introduced M. salmoides, which occupied the majority of the mainstem and several tributaries downstream of barriers to upstream movement. Canonical correspondence analysis showed that only one native species, the Marico barb Barbus motebensis, had a negative spatial association with M. salmoides. Assessment of relative distributions showed this species to be excluded from M. salmoides-invaded river reaches, whereas the other native species were not visibly affected by the invader. This species-specificity of the impact of M. salmoides indicates that their impacts in South African streams may be dependent on predator-naiveté of prey.     

The role of domestic herbivores in endozoochorous plant dispersal in the arid Knersvlakte,South Africa

Herbivores can act as dispersal vectors by purposely or accidentally ingesting ripe fruits, and thus endozoochory is one determining factor for plant distribution patterns. The objective of our study was to investigate to what extent plants of major taxonomic groups of the Knersvlakte (Succulent Karoo, South Africa) are endozoochorously dispersed. On three different farms in the central Knersvlakte, dung of domestic herbivores was collected and analysed by the seedling-emergence method. The resulting species composition was compared to the standing vegetation of thirty-four 1000-m² plots each recorded on one hundred 400-cm² subplots. Our results show that domestic livestock facilitated the dispersal of taxa characteristic of the Knersvlakte, in particular Aizoaceae. Among the taxa of this family, the local endemic dwarf shrub Drosanthemum schoenlandianum emerged with the highest frequency in dung (14.5% of all seedlings). For the Asteraceae, which are frequent in the standing vegetation of the Knersvlakte, however, endozoochorous dispersal by livestock was only of minor importance. Conservation planning should consider these dispersal patterns on behalf of future population dynamics. The complete exclusion of livestock might change current processes and thus alter vegetation patterns.