Big Baby, Little Mother: Tsetse Flies Are Exceptions to the Juvenile Small Size Principle.

While across the animal kingdom offspring are born smaller than their parents, notable exceptions exist. Several dipteran species belonging to the Hippoboscoidea superfamily can produce offspring larger than themselves. In this essay, the blood-feeding tsetse is focused on. It is suggested that the extreme reproductive strategy of this fly is enabled by feeding solely on highly nutritious blood, and producing larval offspring that are soft and malleable. This immense reproductive expenditure may have evolved to avoid competition with other biting flies. Tsetse also transmit blood-borne parasites that cause the fatal diseases called African trypanosomiases. It is discussed how tsetse life history and reproductive strategy profoundly influence the type of vector control interventions used to reduce fly populations. In closing, it is argued that the unusual life history of tsetse warrants their preservation in the areas where human and animal health is not threatened.

"Born to Run"? Not Necessarily: Species and Trait Bias in Persistent Free-Living Transgenic Plants.

The possibility of transgenes from engineered plants ending up in unmanaged populations with undesirable consequences has been a long-term biosafety concern. Experience with traditionally improved plants reveals that most cases of such gene escape have been of little consequence, but on occasion they have led to the evolution of problematic plants or have resulted in an increased extinction risk for wild taxa. Three decades have passed since the first environmental release of transgenic plants, and more than two decades since their first commercialization. Examples of transgenes gone astray are increasingly commonplace. Transgenic individuals have been identified in more than a thousand free-living plant populations. Here I review 14 well-documented consolidated "cases" in which transgenes have found their way into free-living plant populations. Some as transient volunteers; others appear to be persistent transgenic populations. The species involved in the latter are not representative of the current commercialized transgenic crops as whole. They tend to share certain traits that are absent or rare in the transgenic crops that do not exist as persistent populations. The traits commonly occurring in species with persistent transgenic free-living populations are the following, in descending order of importance: (1) a history of occurring as non-transgenic free-living plants, (2) fruits fully or partially shattering prior to harvest, (3) have small or otherwise easily dispersed seeds, either spontaneously or by seed spillage along the supply chain from harvest to consumer, (4) ability to disperse viable pollen, especially to a kilometer or more, (5) perennial habit, and (6) the transgene's fitness effects in the recipient environment are beneficial or neutral. Based on these observations, a thought experiment posits which species might be the next to be reported to occur as free-living transgenic populations.

Does introgression of crop alleles into wild and weedy living populations create cryptic in situ germplasm banks?

Natural gene flow often delivers alleles from cultivated species into unmanaged populations of weedy or wild relatives. Unmanaged populations with introgressed crop alleles may become the unintended repositories of crop alleles that are no longer available to breeders. Descriptive studies have confirmed the introgression of alleles from 17 crop species into free-living populations. Multigeneration experimental studies allowing hybrid-derived individuals to evolve under natural conditions are informative, but remain few. A study in this issue of Molecular Ecology (Molecular Ecology, 26, 2017) documents genomewide crop allele evolution in experimental populations of crop-wild hybrid-derived sunflowers. Approximately, three-quarters of the wild alleles increased in frequency;-however, 5% of the crop alleles in each of the two sites increased (Molecular Ecology, 26, 2017; Figure ). The remainder behaved as if they were neutral. This study's results plus those of prior studies demonstrate that introgressed populations of crop-wild relatives contain a mixture of wild and crop alleles and thereby can be valued as a distinct kind of in situ germplasm resource relative to nonintrogressed populations. [Figure: see text].

Fitness correlates of crop transgene flow into weedy populations: a case study of weedy rice in China and other examples.

Whether transgene flow from crops to cross-compatible weedy relatives will result in negative environmental consequences has been the topic of discussion for decades. An important component of environmental risk assessment depends on whether an introgressed transgene is associated with a fitness change in weedy populations. Several crop-weed pairs have received experimental attention. Perhaps, the most worrisome example is transgene flow from genetically engineered cultivated rice, a staple for billions globally, to its conspecific weed, weedy rice. China's cultivated/weedy rice system is one of the best experimentally studied systems under field conditions for assessing how the presence of transgenes alters the weed's fitness and the likely impacts of that fitness change. Here, we present the cultivated/weedy rice system as a case study on the consequences of introgressed transgenes in unmanaged populations. The experimental work on this system reveals considerable variation in fitness outcomes - increased, decreased, and none - based on the transgenic trait, its introgressed genomic background, and the environment. A review of similar research from a sample of other crop-wild pairs suggests such variation is the rule. We conclude such variation in fitness correlates supports the case-by-case method of biosafety regulation is sound.

Genetic structure reveals a history of multiple independent origins followed by admixture in the allopolyploid weed Salsola ryanii.

It has recently become clear that many invasive species have evolved in situ via hybridization or polyploidy from progenitors which themselves are introduced species. For species formed by hybridization or polyploidy, genetic diversity within the newly formed species is influenced by the number of independent evolutionary origins of the species. For recently formed species, an analysis of genetic structure can provide insight into the number of independent origin events involved in the formation of the species. For a putative invasive allopolyploid species, the number of origins involved in the species formation, the genetic diversity present within these origins, and the level of gene flow between independent origins determines the genetic composition of the neospecies. Here we analyze the genetic structure of the newly formed allopolyploid species, Salsola ryanii, a tumbleweed which evolved within the last 20-100 years in California. We utilize the genetic structure analysis to determine that this new species is the result of at least three independent allopolyplodization events followed by gene flow between the descendants of independent origins.

Maize Germplasm Conservation in Southern California's Urban Gardens: Introduced Diversity Beyond ex situ and in situ Management.

Contemporary germplasm conservation studies largely focus on ex situ and in situ management of diversity within centers of genetic diversity. Transnational migrants who transport and introduce landraces to new locations may catalyze a third type of conservation that combines both approaches. Resulting populations may support reduced diversity as a result of evolutionary forces such as genetic drift, selection, and gene flow, yet they may also be more diverse as a result of multiple introductions, selective breeding and cross pollination among multiple introduced varietals. In this study, we measured the amount and structure of maize molecular genetic diversity in samples collected from home gardens and community gardens maintained by immigrant farmers in Southern California. We used the same markers to measure the genetic diversity and structure of commercially available maize varieties and compared our data to previously reported genetic diversity statistics of Mesoamerican landraces. Our results reveal that transnational dispersal creates an opportunity for the maintenance of maize genetic diversity beyond its recognized centers of diversity.

Rapid range expansion of a newly formed allopolyploid weed in the genus Salsola.

PREMISE OF THE STUDY: Newly formed species (neospecies) can experience a variety of demographic fates, ranging from rapid invasive expansion to rapid extinction. Here we investigate the fate of the neospecies Salsola ryanii 10 years after its initial discovery in the Central Valley of California, USA. This species is an allopolyploid derived via hybridization between the invasive species, S. australis and S. tragus. METHODS: We conducted a systematic collection of Salsola species from 53 sites in California. Species-specific intersimple sequence repeat (ISSR) markers were used to determine the species of each individual collected. The range of S. ryanii identified in this study was compared to the range in 2002 to determine how the range has shifted in the decade between surveys. KEY RESULTS: In this survey, we identified 15 sites where S. ryanii was present (28% of sites), a significant population number increase since 2002. CONCLUSIONS: Salsola ryanii has undergone a dramatic population number expansion in the decade since it was originally documented. We are not aware of any plant neospecies whose range spontaneously experienced such a dramatic expansion. Salsola ryanii has every indication of being just as invasive as its highly invasive parents.

Risk assessment of gene flow from genetically engineered virus resistant cassava to wild relatives in Africa: an expert panel report.

The probability and consequences of gene flow to wild relatives is typically considered in the environmental risk assessment of genetically engineered crops. This is a report from a discussion by a group of experts who used a problem formulation approach to consider existing information for risk assessment of gene flow from cassava (Manihot esculenta) genetically engineered for virus resistance to the 'wild' (naturalized) relative M. glaziovii in East Africa. Two environmental harms were considered in this case: (1) loss of genetic diversity in the germplasm pool, and (2) loss of valued species, ecosystem resources, or crop yield and quality due to weediness or invasiveness of wild relatives. Based on existing information, it was concluded that gene flow will occur, but it is not likely that this will reduce the genetic diversity in the germplasm pool. There is little existing information about the impact of the virus in natural populations that could be used to inform a prediction about whether virus resistance would lead to an increase in reproduction or survival, hence abundance of M. glaziovii. However, an increase in the abundance of M. glaziovii should be manageable, and would not necessarily lead to the identified environmental harms.

Novel seed protection in the recently evolved invasive, California wild radish, a hybrid Raphanus sp. (Brassicaceae).

UNLABELLED: • PREMISE OF THE STUDY: Interspecific hybridization may have considerable effects on plant structural defenses that can contribute to the success of invasive hybrid lineages. Changes in fruit structural and material properties are predicted to have key effects on predispersal granivory.• METHODS: Here, we asked whether plant structure can increase the fitness of a hybrid invasive relative to its progenitors. We compared fruit traits of the hybrid-derived lineage, California wild radish, with its progenitors, cultivated radish and jointed charlock.• KEY RESULTS: The hybrid lineage is significantly different from one or both ancestors in fruit length, mass, diameter, volume, shape, wall strength, and internal seed distribution. We experimentally exposed the fruits of both hybrid and wild progenitor to avian granivores and found (1) different types and degrees of damage at the different fruit sections and (2) significant differences in the inflicted damage at different sections of the fruit.• CONCLUSIONS: Combining our descriptive and experimental data, we conclude that the novel seed protection of the hybrid California wild radish is an important defense mechanism. It offers differential protection to its seeds and according to our findings, better protection of seeds that have been found to be better competitors. We suggest then that the fruit has enabled, at least in part, the successful replacement of the parental species by the hybrid lineage.

Is gene flow the most important evolutionary force in plants?

Although theory has demonstrated rather low levels of gene flow are sufficient to counteract opposing mutation, drift, and selection, widespread recognition of the evolutionary importance of gene flow has come slowly. The perceived role of gene flow as an evolutionary force has vacillated over the last century. In the last few decades, new methods and analyses have demonstrated that plant gene flow rates vary tremendously-from nil to very high-depending on the species and specific populations involved, and sometimes over time for individual populations. In many cases, the measured gene flow rates are evolutionarily significant at distances of hundreds and sometimes thousands of meters, occurring at levels sufficient to counteract drift, spread advantageous alleles, or thwart moderate levels of opposing local selection. Gene flow in plants is likely to often act as a cohesive force, uniting individual plant species into real evolutionary units. Also, gene flow can evolve under natural selection, decreasing or increasing. The fact of frequent, but variable, plant gene flow has important consequences for applied issues in which the presence or absence of gene flow might influence the outcome of a policy, regulatory, or management decision. Examples include the unintended spread of engineered genes, the evolution of invasiveness, and conservation. New data-rich genomic techniques allow closer scrutiny of the role of gene flow in plant evolution. Most plant evolutionists now recognize the importance of gene flow, and it is receiving increased recognition from other areas of plant biology as well.

Taxonomic and life history bias in herbicide resistant weeds: implications for deployment of resistant crops.

Evolved herbicide resistance (EHR) is an important agronomic problem and consequently a food security problem, as it jeopardizes herbicide effectiveness and increases the difficulty and cost of weed management. EHR in weeds was first reported in 1970 and the number of cases has accelerated dramatically over the last two decades. Despite 40 years of research on EHR, why some weeds evolve resistance and others do not is poorly understood. Here we ask whether weed species that have EHR are different from weeds in general. Comparing taxonomic and life history traits of weeds with EHR to a control group ("the world's worst weeds"), we found weeds with EHR significantly over-represented in certain plant families and having certain life history biases. In particular, resistance is overrepresented in Amaranthaceae, Brassicaceae and Poaceae relative to all weeds, and annuality is ca. 1.5 times as frequent in weeds with EHR as in the control group. Also, for perennial EHR weeds, vegetative reproduction is only 60% as frequent as in the control group. We found the same trends for subsets of weeds with EHR to acetolactate synthase (ALS), photosystem II (PSII), and 5-enolpyruvylshikimate-3-phosphate (EPSP) synthase-inhibitor herbicides and with multiple resistance. As herbicide resistant crops (transgenic or not) are increasingly deployed in developing countries, the problems of EHR could increase in those countries as it has in the USA if the selecting herbicides are heavily applied and appropriate management strategies are not employed. Given our analysis, we make some predictions about additional species that might evolve resistance.

The genomic signature of crop-wild introgression in maize.

The evolutionary significance of hybridization and subsequent introgression has long been appreciated, but evaluation of the genome-wide effects of these phenomena has only recently become possible. Crop-wild study systems represent ideal opportunities to examine evolution through hybridization. For example, maize and the conspecific wild teosinte Zea mays ssp. mexicana (hereafter, mexicana) are known to hybridize in the fields of highland Mexico. Despite widespread evidence of gene flow, maize and mexicana maintain distinct morphologies and have done so in sympatry for thousands of years. Neither the genomic extent nor the evolutionary importance of introgression between these taxa is understood. In this study we assessed patterns of genome-wide introgression based on 39,029 single nucleotide polymorphisms genotyped in 189 individuals from nine sympatric maize-mexicana populations and reference allopatric populations. While portions of the maize and mexicana genomes appeared resistant to introgression (notably near known cross-incompatibility and domestication loci), we detected widespread evidence for introgression in both directions of gene flow. Through further characterization of these genomic regions and preliminary growth chamber experiments, we found evidence suggestive of the incorporation of adaptive mexicana alleles into maize during its expansion to the highlands of central Mexico. In contrast, very little evidence was found for adaptive introgression from maize to mexicana. The methods we have applied here can be replicated widely, and such analyses have the potential to greatly inform our understanding of evolution through introgressive hybridization. Crop species, due to their exceptional genomic resources and frequent histories of spread into sympatry with relatives, should be particularly influential in these studies.

Sequence polymorphisms in wild, weedy, and cultivated rice suggest seed-shattering locus sh4 played a minor role in Asian rice domestication.

The predominant view regarding Asian rice domestication is that the initial origin of nonshattering involved a single gene of large effect, specifically, the sh4 locus via the evolutionary replacement of a dominant allele for shattering with a recessive allele for reduced shattering. Data have accumulated to challenge this hypothesis. Specifically, a few studies have reported occasional seed-shattering plants from populations of the wild progenitor of cultivated rice (Oryza rufipogon complex) being homozygous for the putative "nonshattering" sh4 alleles. We tested the sh4 hypothesis for the domestication of cultivated rice by obtaining genotypes and phenotypes for a diverse set of samples of wild, weedy, and cultivated rice accessions. The cultivars were fixed for the putative "nonshattering" allele and nonshattering phenotype, but wild rice accessions are highly polymorphic for the putative "nonshattering" allele (frequency ∼26%) with shattering phenotype. All weedy rice accessions are the "nonshattering" genotype at the sh4 locus but with shattering phenotype. These data challenge the widely accepted hypothesis that a single nucleotide mutation ("G"/"T") of the sh4 locus is the major driving force for rice domestication. Instead, we hypothesize that unidentified shattering loci are responsible for the initial domestication of cultivated rice through reduced seed shattering.

Reproductive isolation during domestication.

It has been hypothesized that reproductive isolation should facilitate evolution under domestication. However, a systematic comparison of reproductive barrier strength between crops and their progenitors has not been conducted to test this hypothesis. Here, we present a systematic survey of reproductive barriers between 32 economically important crop species and their progenitors to better understand the role of reproductive isolation during the domestication process. We took a conservative approach, avoiding those types of reproductive isolation that are poorly known for these taxa (e.g., differences in flowering time). We show that the majority of crops surveyed are isolated from their progenitors by one or more reproductive barriers, despite the fact that the most important reproductive barrier in natural systems, geographical isolation, was absent, at least in the initial stages of domestication for most species. Thus, barriers to reproduction between crops and wild relatives are closely associated with domestication and may facilitate it, thereby raising the question whether reproductive isolation could be viewed as a long-overlooked "domestication trait." Some of the reproductive barriers observed (e.g., polyploidy and uniparental reproduction), however, may have been favored for reasons other than, or in addition to, their effects on gene flow.

Maize x Teosinte Hybrid Cobs Do Not Prevent Crop Gene Introgression.

Maize x Teosinte Hybrid Cobs Do Not Prevent Crop Gene Introgression. Whether introgression from crops to wild relatives can occur is an important component of transgene risk assessment. In the case of maize, which co-occurs with its wild relative teosinte in Mexico, the possibility of introgression has been controversial. Maize is cross-compatible with teosinte, and spontaneous hybridization is known to occur. Some scientists have hypothesized that the maize x teosinte cob infructescence will prevent progeny dispersal, thus preventing introgression. Motivated by a prior study where we found maize x teosinte hybrid fruits naturally dispersed under field conditions, we tested whether hybrid cobs hold their fruits as tightly as maize cobs. We found the force required to detach hybrid fruits was substantially and significantly less than that for maize. Consequently, we expect that introgression of transgenes from maize into teosinte in Mexico should occur largely unimpeded by the hybrid cob.La mazorca o elote híbrido de maíz x teocintle no impide la introgresión de genes transgénicos provenientes del cultivo. La introgresión entre el maíz cultivado y el maíz silvestre, o teocintle, es un componente importante en la evaluación ambiental relacionada con los riesgos de la introducción de genes transgénicos. La posibilidad de introgresión entre el maíz domesticado y el teocintle ha sido un tema controversial, en particular en México, donde maíz y teocintle coexisten. El maíz es compatible con el teocintle y la hibridización espontánea ocurre entre ellos. Algunos científicos han planteado como hipótesis que al cruzar el maíz con teocintle, la estructura interna de la infrutescencia que sujeta los frutos conocida como la mazorca de maíz o el elote, impide la dispersión de la progenie evitando que la introgresión ocurra. Los resultados de un estudio previo evidencian la dispersión de los frutos híbridos del maíz x teocintle en condiciones naturales. Motivados por estos resultados, hemos decidido investigar si la mazorca o el elote de las infrutescencias del híbrido sujetan los frutos con una fuerza comparable o mayor a la del maíz. Nuestras mediciones implican que la fuerza necesaria para liberar los frutos híbridos son substancial y significativamente menores que aquellas necesarias para desprender los frutos del maíz. Como conclusión sugerimos que en México, la mazorca o el elote no representan una barrera que impida la introgresión de los genes transgénicos del maíz al teocintle.

Rapid evolutionary divergence and ecotypic diversification of germination behavior in weedy rice populations.

Feral plants have evolved from well-studied crops, providing good systems for elucidation of how weediness evolves. As yet, they have been largely neglected for this purpose. The evolution of weediness can occur by simple back mutations in domestication genes (domestication in reverse). Whether the evolutionary steps to weediness always occur in reverse remains largely unknown. We examined seed germination behavior in recently evolved weedy rice (Oryza sativa f. spontanea) populations and their coexisting cultivars in eastern and north-eastern China to address whether 'dedomestication' is the simple reverse of domestication. We found that these weedy populations did not diverge from their progenitors by reverting to the pre-domestication trait of seed dormancy. Instead, they have evolved a novel mechanism to avoid growing in inappropriate environments via changes in critical temperature cues for seed germination. Furthermore, we found evidence for subsequent ecotypic divergence of these populations such that the critical temperature for germination correlates with the local habitat temperature at latitudinal gradients. The origins of problematic plant species, weeds and invasives, have already been studied in detail. These plants can thus be used as systems for studying rapid evolution. To determine whether and how that evolution is adaptive, experiments such as those described here can be performed.

Rapid evolution of morphology and adaptive life history in the invasive California wild radish (Raphanus sativus) and the implications for management.

Understanding the evolution and demography of invasive populations may be key for successful management. In this study, we test whether or not populations of the non-native, hybrid-derived California wild radish have regionally adapted to divergent climates over their 150-year history in California and determine if population demographic dynamics might warrant different region-specific strategies for control. Using a reciprocal transplant approach, we found evidence for genetically based differences both between and among northern, coastal and southern, inland populations of wild radish. Individual fitness was analyzed using a relatively new statistical method called 'aster modeling' which integrates temporally sequential fitness measurements. In their respective home environments, fitness differences strongly favored southern populations and only slightly favored northern populations. Demographic rates of transition and sensitivities also differed between regions of origin, suggesting that the most effective approach for reducing overall population growth rate would be to target different life-history stages in each region.