Stephen M. Hovick

Does phylogeny matter? Assessing the impact of phylogenetic information in ecological meta‐analysis

Meta-analysis is increasingly used in ecology and evolutionary biology. Yet, in these fields this technique has an important limitation: phylogenetic non-independence exists among taxa, violating the statistical assumptions underlying traditional meta-analytic models. Recently, meta-analytical techniques incorporating phylogenetic information have been developed to address this issue. However, no syntheses have evaluated how often including phylogenetic information changes meta-analytic results. To address this gap, we built phylogenies for and re-analysed 30 published meta-analyses, comparing results for traditional vs. phylogenetic approaches and assessing which characteristics of phylogenies best explained changes in meta-analytic results and relative model fit. Accounting for phylogeny significantly changed estimates of the overall pooled effect size in 47% of datasets for fixed-effects analyses and 7% of datasets for random-effects analyses. Accounting for phylogeny also changed whether those effect sizes were significantly different from zero in 23 and 40% of our datasets (for fixed- and random-effects models, respectively). Across datasets, decreases in pooled effect size magnitudes after incorporating phylogenetic information were associated with larger phylogenies and those with stronger phylogenetic signal. We conclude that incorporating phylogenetic information in ecological meta-analyses is important, and we provide practical recommendations for doing so.

The scope of Baker's law

Bakers law refers to the tendency for species that establish on islands by long-distance dispersal to show an increased capacity for self-fertilization because of the advantage of self-compatibility when colonizing new habitat. Despite its intuitive appeal and broad empirical support, it has received substantial criticism over the years since it was proclaimed in the 1950s, not least because it seemed to be contradicted by the high frequency of dioecy on islands. Recent theoretical work has again questioned the generality and scope of Bakers law. Here, we attempt to discern where the idea is useful to apply and where it is not. We conclude that several of the perceived problems with Bakers law fall away when a narrower perspective is adopted on how it should be circumscribed. We emphasize that Bakers law should be read in terms of an enrichment of a capacity for uniparental reproduction in colonizing situations, rather than of high selfing rates. We suggest that Bakers law might be tested in four different contexts, which set the breadth of its scope: the colonization of oceanic islands, metapopulation dynamics with recurrent colonization, range expansions with recurrent colonization, and colonization through species invasions.

Hybridisation is associated with increased fecundity and size in invasive taxa: meta-analytic support for the hybridisation-invasion hypothesis.

The hypothesis that interspecific hybridisation promotes invasiveness has received much recent attention, but tests of the hypothesis can suffer from important limitations. Here, we provide the first systematic review of studies experimentally testing the hybridisation-invasion (H-I) hypothesis in plants, animals and fungi. We identified 72 hybrid systems for which hybridisation has been putatively associated with invasiveness, weediness or range expansion. Within this group, 15 systems (comprising 34 studies) experimentally tested performance of hybrids vs. their parental species and met our other criteria. Both phylogenetic and non-phylogenetic meta-analyses demonstrated that wild hybrids were significantly more fecund and larger than their parental taxa, but did not differ in survival. Resynthesised hybrids (which typically represent earlier generations than do wild hybrids) did not consistently differ from parental species in fecundity, survival or size. Using meta-regression, we found that fecundity increased (but survival decreased) with generation in resynthesised hybrids, suggesting that natural selection can play an important role in shaping hybrid performance – and thus invasiveness – over time. We conclude that the available evidence supports the H-I hypothesis, with the caveat that our results are clearly driven by tests in plants, which are more numerous than tests in animals and fungi.

Evaluating the post-release efficacy of invasive plant biocontrol by insects: a comprehensive approach

We propose a comprehensive program to evaluate the post-release phase of biocontrol programs that use insect herbivores to control invasive plant species. We argue that any release should be done in randomized release and non-release sites and should be followed up by well-replicated sampling and experimental protocols that evaluate the degree of success or failure. These follow-up studies should include landscape scale monitoring across relevant habitat gradients of (1) the abundance of the biocontrol agent, (2) the impact of the biocontrol agent on the target plant species, (3) the potential for non-target effects, and (4) the response of native species and communities to a reduction in the invasive species. We also argue that (5) experimental reductions of the biocontrol agent are required to eliminate the chance that the putative impact of the biocontrol agent is not confounded with other causes. Finally, we describe six scenarios, informed largely by a community ecology perspective, in which a biocontrol agent may decrease the abundance or vigor of the target plant species but not lead to successful control where native communities re-establish. We classify these failure scenarios as either direct or indirect effects of the invasive plant species: Native Source Limitation, Static Competitive Hierarchies, Novel Weapons, Trophic Shifts, Invasive Engineering and Associated Invasives. Overall, we argue that well replicated and landscape-scale post release monitoring programs are required not only to evaluate critically the degree of success and failure of biocontrol programs worldwide but also to provide insights into improving future biocontrol efforts.

Hybridization Alters Early Life-History Traits and Increases Plant Colonization Success in a Novel Region

Hybridization is hypothesized to promote invasiveness, but empirical tests comparing the performance of hybrid taxa versus parental taxa in novel regions are lacking. We experimentally compared colonization ability of populations of wild radish (Raphanus raphanistrum) with populations of advanced-generation hybrids between wild radish and cultivated radish (Raphanus sativus) in a southeast Texas pasture, well beyond the known invasive range of hybrid radish. We also manipulated the strength of interspecific competition to better generalize across variable environments. In both competitive environments, hybrid populations produced at least three times more seeds than did wild radish populations, a distinction that was driven by greater hybrid seedling emergence, earlier hybrid emergence, and more hybrid seedlings surviving to flower, rather than by greater individual fecundity. Flowering duration in hybrids was less negatively affected by competition than it was in wild radish, while early emergence was associated with subsequent high seed output in both biotypes. Our data show that hybridization can enhance colonization success in a novel region and, by comparison with previous studies, that the life-history traits enhancing hybrid success can differ across regions, even for lineages originating from the same hybridization event. These results imply a much larger arena for hybrid success than previously appreciated.

Quantitative trait locus mapping identifies candidate alleles involved in adaptive introgression and range expansion in a wild sunflower

The wild North American sunflowers Helianthus annuus and H. debilis are participants in one of the earliest identified examples of adaptive trait introgression, and the exchange is hypothesized to have triggered a range expansion in H. annuus. However, the genetic basis of the adaptive exchange has not been examined. Here, we combine quantitative trait locus (QTL) mapping with field measurements of fitness to identify candidate H. debilis QTL alleles likely to have introgressed into H. annuus to form the natural hybrid lineage H. a. texanus. Two 500‐individual BC1 mapping populations were grown in central Texas, genotyped for 384 single nucleotide polymorphism (SNP) markers and then phenotyped in the field for two fitness and 22 herbivore resistance, ecophysiological, phenological and architectural traits. We identified a total of 110 QTL, including at least one QTL for 22 of the 24 traits. Over 75% of traits exhibited at least one H. debilis QTL allele that would shift the trait in the direction of the wild hybrid H. a. texanus. We identified three chromosomal regions where H. debilis alleles increased both female and male components of fitness; these regions are expected to be strongly favoured in the wild. QTL for a number of other ecophysiological, phenological and architectural traits colocalized with these three regions and are candidates for the actual traits driving adaptive shifts. G × E interactions played a modest role, with 17% of the QTL showing potentially divergent phenotypic effects between the two field sites. The candidate adaptive chromosomal regions identified here serve as explicit hypotheses for how the genetic architecture of the hybrid lineage came into existence.

Tailoring biocontrol to maximize top-down effects: on the importance of underlying site fertility.

The degree to which biocontrol agents impact invasive plants varies widely across landscapes, often for unknown reasons. Understanding this variability can help optimize invasive species management while also informing our understanding of trophic linkages. To address these issues, we tested three hypotheses with contrasting predictions regarding the likelihood of biocontrol success. (1) The biocontrol effort hypothesis: invasive populations are regulated primarily by top-down effects, predicting that increased biocontrol efforts alone (e.g., more individuals of a given biocontrol agent or more time since agent release) will enhance biocontrol success. (2) The relative fertility hypothesis: invasive populations are regulated primarily by bottom-up effects, predicting that nutrient enrichment will increase dominance by invasives and thus reduce biocontrol success, regardless of biocontrol efforts. (3) The fertility-dependent biocontrol effort hypothesis: top-down effects will only regulate invasive populations if bottom-up effects are weak. It predicts that greater biocontrol efforts will increase biocontrol success, but only in low-nutrient sites. To test these hypotheses, we surveyed 46 sites across three states with prior releases of Galerucella beetles, the most common biocontrol agents used against invasive purple loosestrife (Lythrum salicaria). We found strong support for the fertility-dependent biocontrol effort hypothesis, as biocontrol success occurred most often with greater biocontrol efforts, but only in low-fertility sites. This result held for early stage metrics of biocontrol success (higher Galerucella abundance) and ultimate biocontrol outcomes (decreased loosestrife plant size and abundance). Presence of the invasive grass Phalaris arundinacea was also inversely related to loosestrife abundance, suggesting that biocontrol-based reductions in loosestrife made secondary invasion by P. arundinacea more likely. Our data suggest that low-nutrient sites be prioritized for loosestrife biocontrol and that future monitoring account for variation in site fertility or work to mitigate it. We introduce a new framework that integrates our findings with conflicting patterns previously reported from other biocontrol systems, proposing a unimodal relationship whereby nutrient availability enhances biocontrol success in low-nutrient sites but hampers it in high-nutrient sites. Our results represent one of the first examples of biocontrol success depending on site fertility, which has the potential to inform biocontrol-based management decisions across entire regions and among contrasting systems.

Community dominance patterns, not colonizer genetic diversity, drive colonization success in a test using grassland species

Successful colonization and/or invasion depend on characteristics of the invaded community and of the colonizer itself. Although many studies have documented a negative relationship between invasibility and biodiversity, the importance of community evenness is rarely examined and thus poorly understood. However, colonizer characteristics, including population genetic diversity, can also be important determinants of colonization success. We conducted a greenhouse experiment to assess the relative importance of community evenness and colonizer population genetic diversity using the weed Arabidopsis thaliana. We added seeds of A. thaliana (varying genetic diversity while keeping propagule pressure constant) to four types of constructed plant communities: those dominated by legumes, grasses or forbs, or with equal abundances of all three functional groups. We selected community members from a large pool of species to avoid the confounding effects of species identity. We also assessed the success of multiple seedbank colonizers to assess generality in the effects of our evenness treatments. Equal-abundance communities were no better at suppressing colonization than communities dominated by a single functional group. Forb-dominated communities suppressed A. thaliana colonization better than grass-dominated communities and suppressed seedbank colonizers better than legume-dominated communities. Equal-abundance communities were similar to forb-dominated ones in their eventual composition and in their invasibility, suggesting that forbs drove colonizer suppression in that treatment rather than high evenness itself. Most of our forbs grew quickly, yielding productive forb-dominated communities; this points to the importance of growth and colonization phenology in our system. A. thaliana genetic diversity did not affect colonization success, perhaps because strong interspecific competition substantially limited A. thaliana seedling emergence.

Funding needed for assessments of weed biological control

Invasive non-native plants are a serious economic and ecological problem worldwide, and major efforts are therefore devoted to reducing weed abundance in agricultural and natural settings. Effective options for reducing invasive abundance and spread are few, although one common approach is biological control - the introduction of specialist herbivores or pathogens from a weeds native range to suppress weed abundance in the introduced range. Biocontrol is a crucial tool in invasive species management because, once biocontrol agents establish, they are often self-sustaining and can greatly reduce invasive populations.

A reassessment of the genome size–invasiveness relationship in reed canarygrass (Phalaris arundinacea)

Background and Aims Genome size is hypothesized to affect invasiveness in plants. Key evidence comes from a previous study of invasive eastern North American populations of the grass Phalaris arundinacea: invasive genotypes with smaller genomes had higher growth rates, and genome sizes were smaller in the invasive vs. native range. This study aimed to re-investigate those patterns by examining a broader range of North American populations and by employing the modern best-practice protocol for plant genome size estimation in addition to the previously used protocol. Methods Genome sizes were measured using both internal and pseudo-internal standardization protocols for 20 invasive and nine native range accessions of P. arundinacea. After a round of vegetative propagation to reduce maternal environmental effects, growth (stem elongation) rates of these accessions were measured in the greenhouse. Key Results Using the best-practice protocol, there was no evidence of a correlation between genome size and growth rates (P = 0.704), and no evidence for differences in genome sizes of invasive and native range accessions (P > 0.353). However, using the older genome size estimation protocol, both relationships were significant (reproducing the results of the previous study). Conclusions Genome size reduction has not driven increased invasiveness in a broad sample of North American P. arundinacea. Further, inappropriate genome size estimation techniques can create spurious correlations between genome size and plant traits such as growth rate. Valid estimation is vital to progress in understanding the potentially widespread effects of genome size on biological processes and patterns.