Highly diverse and highly successful: invasive Australian acacias have not experienced genetic bottlenecks globally

Background and AimsInvasive species may undergo rapid evolution despite very limited standing genetic diversity. This so-called genetic paradox of biological invasions assumes that an invasive species has experienced (and survived) a genetic bottleneck and then underwent local adaptation in the new range. In this study, we test how often Australian acacias (genus Acacia), one of the world’s most problematic invasive tree groups, have experienced genetic bottlenecks and inbreeding.MethodsWe collated genetic data from 51 different genetic studies on Acacia species to compare genetic diversity between native and invasive populations. These studies analysed 37 different Acacia species, with genetic data from the invasive ranges of 11 species, and data from the native range for 36 species (14 of these 36 species are known to be invasive somewhere in the world, and the other 22 are not known to be invasive).Key ResultsLevels of genetic diversity are similar in native and invasive populations, and there is little evidence of invasive populations being extensively inbred. Levels of genetic diversity in native range populations also did not differ significantly between species that have and that do not have invasive populations.ConclusionWe attribute our findings to the impressive movement, introduction effort and human usage of Australian acacias around the world.     

Two alien invasive acacias in Italy: Differences and similarities in their flowering and insect visitors

In Italy, alien acacias have been introduced for ornamental and reforestation purposes, and some species became invasive occupying patches of the Mediterranean landscape. On the Island of Elba (Central Italy), Acacia dealbata and A. pycnantha form dense stands at short distance, showing an impressive massive flowering at the end of the winter/early spring. Our aim was to investigate the behaviour of the two species in relation to the flowering features, from phenology to floral characteristics, and their replay to the observed flower visitors. Differences between the two species emerged on all the parameters considered. A. pycnantha peak of flowering occurred later than A. dealbata and showed larger flower heads (FHs), more flowers/head, stamens/flower and polyads. On A. dealbata, we recorded longer racemes and more FHs/raceme, determining a more flower-dense crown. Even if contacts with flower visitors were generally low on both species, A. dealbata showed a more heterogeneous visitor assemblage. Both acacias species interacted with local generalist pollinators, as bumblebees and honey bees. Flower handling and resource collection strategy by the honey bee indicate a long-term relationship between the bee and the acacias, with bees investing longer time on the larger A. pycnantha flower heads.     

西藏高原高寒植物长鞭红景天生殖特征成功及影响因子

为了探索高寒植物长鞭红景天(Rhodiola fastigiata)濒危的生殖生态学机理,寻找保护和恢复该物种的有效途径,对其繁育系统、开花物候特征、生殖过程(开花、结实、种子产量和发育)及其影响生殖成功的主要因子进行调查分析。结果表明:长鞭红景天包括两性植株和雌性植株两种类型,其中两性植株仅执行雄性功能,种群繁殖为异交,属风媒传粉方式。种群花期出现在1a中降雨和气温较高的季节,6月下旬开始开花,7月下旬结束,时间30d左右。单株开花动态呈现单峰和双峰曲线两种模式,两性植株花期(9.29d)大于雌性植株花期(7.06d),并先于雌性植株开放,但雌性植株开花的高峰期比两性花早。从花蕾到开花,雌性单株有16%—25%的花蕾损失;开花到结实,单株36%—57%的花败育;至种子成熟,共有84%—94%的胚珠损失,仅6%—16%的胚珠发育成种子。雌性单株平均种子产量估计值约为285粒,说明长鞭红景天生殖潜力较大。人工条件下种子发芽率为93.6%,可见种子质量不是导致其濒危的原因,花粉限制和种子产量可能是影响生殖成功的主要因素。     

Sprague's pipit breeding biology and reproductive success in planted and native grasslands

Much of the native grasslands in agricultural regions have been converted to cropland or tilled and seeded with non‐native grasses for livestock production. Several grassland songbird species occupy planted grasslands, but occupancy or density may not be a reliable indicator of habitat quality. I studied the breeding biology of Sprague's pipit Anthus spragueii from 2004 to 2008 in Saskatchewan, Canada. My objective was to determine the extent to which the breeding biology, density and reproductive success of pipits varied in planted and native grasslands. Peak clutch initiation occurred in mid‐ to late‐May in planted and native grassland. Peak pipit density also occurred in May, but density drastically declined over the breeding season in planted grassland. Clutch size varied among years and declined over the breeding season, but was similar in planted (4.7  0.1 SE) and native grasslands (4.5  0.1 SE). Daily nest survival rates varied with age of the nest and date, but the relationships differed in the two habitats and was likely a result of lower nestling survival in planted grassland compared to native grassland. The number of young fledged per nest increased as the season progressed and tended to be greater in native (1.2  0.1 SE) than planted (0.9  0.2 SE) grasslands. Seasonal productivity was much greater in native grassland. Only three nests were initiated after May in planted grassland and all were unsuccessful, whereas pipit young fledged at higher rates from nests initiated in native grassland in June and July than planted grassland nests initiated in May. The number of fledged young from successful nests did not vary strongly with habitat, date or year. This research indicates that planted grasslands attract pipits at the beginning of the breeding season, but habitat suitability and reproductive success substantially declines as the breeding season progresses compared to that found in native grassland.     

How flower biology and breeding system affect the reproductive success of the narrow endemic Polygala vayredae Costa (Polygalaceae)

Polygala vayredae is a narrow endemic species from the oriental pre-Pyrenees. Despite its conservation status and rarity, no information is available on its reproductive biology. As the flower is the structure directly involved in pollinator attraction, its morphological and functional traits have major effects on the reproductive success of the plant. In this work, the flower biology and breeding system of P. vayredae were studied to evaluate how they affect the reproductive outcome in natural populations. Flower morphology, flower rewards, and male and female functioning throughout the lifespan of the flower were assessed. Pollination experiments, involving pollinator exclusion and pollen from different sources, were conducted, and the pollen ovule index was determined. Female fitness and the occurrence of pollen limitation were assessed in three natural populations over 2 years by observing the presence of pollen on the stigma, pollen tube development, and fruit production. Polygala vayredae flowers are elaborate and long-lived with nectar rewards. The floral traits are well adapted to xenogamy and entomophily, which are in accordance with the observed breeding system and auto-incompatibility system. No mechanism of reproductive assurance was observed and P. vayredae strictly depends on pollinators to set fruit. Low fruit production was observed in the studied populations, which was largely the result of scarce, unreliable, and/or inefficient pollinators and poor pollen quality. In addition, available resources may be a limiting factor. The reproductive strategy of P. vayredae prevents inbreeding depression by a self-incompatibility system, which in years of scarce pollinators is overcome by the plant habit.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 67–81.     

Reproductive biology of Azadirachta indica (Meliaceae), a medicinal tree species from arid zones

Neem or the Margosa tree, Azadirachta indica (Meliaceae), is a versatile tree species common in arid zones. We investigated the reproductive biology of Neem, including phenology, floral biology, pollination ecology and the breeding system, in three natural populations in northern India. Phenological observations established that trees within a population flowered synchronously, but populations varied in the time of onset of flowering; a few trees flowered twice a year. An inflorescence bears ～ 91 bisexual, white and sweet‐scented flowers. The stigma consists of a non‐receptive apical region and a subjacent rim of receptive surface; papillae on the two regions differ in their morphology and function. The occurrence of natural pollination both by wind and insects indicates ambophily. Insect pollinators were predominantly represented by Hymenoptera and Lepidoptera; Apis spp. were the most effective pollinators. Although the occurrence of incomplete dichogamy and a pollen : ovule ratio of ～ 627 indicates the possibility of autogamy, supplemental pollinations clearly established that the trees were 100% self‐incompatible and incompatibility is regulated at the gametophytic level. Natural fruit set was low (～ 5%) and hand cross‐pollinations increased the fruit set to ～ 19%. Our study demonstrated that natural fruit set in Neem is possibly limited by insufficient amounts of xenogamous pollination and resources.     

Reproductive success and helper effects in the cooperatively breeding grey-crowned babbler

Cooperative breeding, where some individuals help to raise offspring that are not their own, is a relatively rare social system in birds. We studied the breeding biology of a declining cooperative breeder, the grey-crowned babbler Pomatostomus temporalis , with the aim of isolating the social factors that affect its reproductive success. Most breeding pairs were assisted by philopatric offspring, although pairs could breed successfully without helpers. Females laid up to four clutches (usually three eggs per clutch) per season. Male (but not female) helpers increased the number of young fledged from individual nests and the likelihood of re-nesting, resulting in higher seasonal fledgling production. Helper effects on brood size and fledgling production were greater in the second year of the study, which was also characterized by higher nest failure. This suggests that helpers enhance reproduction more in poor conditions. Our study demonstrates the interacting effects of social and ecological factors on reproductive success, and that retention of offspring is not always beneficial for the breeders in cooperative species.     

Reproductive ecology of the endangered Alpine species Eryngium alpinum L. (Apiaceae): phenology, gene dispersal and reproductive success

BACKGROUND AND AIMS: Eryngium alpinum (Apiaceae) is an endangered perennial, characteristic of the Alpine flora. Because the breeding system influences both demographic (reproductive success) and genetic (inbreeding depression, evolutionary potential) parameters that are crucial for population maintenance, the reproductive ecology of E. alpinum was investigated. Specifically, the aims of the study were (1) to determine the factors (resources and/or pollen) limiting plant fitness; and (2) to assess the potential for gene flow within a plant, within a patch of plants, and across a whole valley where the species is abundant. METHODS: Field experiments were performed at two sites in the Fournel valley, France, over three consecutive years. Studies included a phenological survey, observations of pollinators (visitation rates and flight distances), dispersal of a fluorescent powder used as a pollen analogue, the use of seed traps, determination of the pollen/ovule ratio, and an experiment to test whether seed production is limited by pollen and/or by resources. KEY RESULTS: E. alpinum is pollinated by generalist pollinators, visitation rates are very high and seed set is resource- rather than pollen-limited. The short flights of honeybees indicate a high potential for geitonogamy, and low pollen and seed dispersals suggest strong genetic structure over short distances. These results are interpreted in the light of previous molecular markers studies, which, in contrast, showed complete outcrossing and high genetic homogeneity. CONCLUSION:S. The study highlights the usefulness of adopting several complementary approaches to understanding the dynamic processes at work in natural populations, and the conservation implications for E. alpinum are emphasized. Although the studied populations do not seem threatened in the near future, long-term monitoring appears necessary to assess the impact of habitat fragmentation. Moreover, this study provides useful baseline data for future investigations in smaller and more isolated populations.     

Migration patterns and changes in population biology associated with the worldwide spread of the oilseed rape pathogen Leptosphaeria maculans

Pathogen introductions into novel areas can lead to the emergence of new fungal diseases of plants. Understanding the origin, introduction pathways, possible changes in reproductive system and population size of fungal pathogens is essential in devising an integrated strategy for the control of these diseases. We used minisatellite markers to infer the worldwide invasion history of the fungal plant pathogen Leptosphaeria maculans, which causes stem canker (blackleg) of oilseed and vegetable brassicas. Clustering analyses partitioned genotypes into distinct populations corresponding to major geographic regions, along with two differentiated populations in Western Canada. Comparison of invasion scenarios using Approximate Bayesian Computation suggested an origin of the pathogen in the USA, the region where epidemics were first recorded, and independent introductions from there over the last few decades into Eastern Canada (Ontario), Europe and Australia. The population in Western Canada appeared to be founded from a source in Ontario and the population in Chile resulted from an admixture between multiple sources. A bottleneck was inferred for the introduction into Western Canada but not into Europe, Ontario or Australia. Clonality appeared high in Western Canada, possibly because environmental conditions there were less conducive to sexual reproduction. Leptosphaeria maculans is a model invasive pathogen with contrasting features in different regions: shallow population structure, high genetic variability and regular sexual recombination in some regions, by comparison with reduced genetic variability, high rates of asexual multiplication, strong population structure or admixture in others.     

Do introduced honeybees (Apis mellifera,Hymenoptera) provide full pollination service to bird‐adapted Australian plants with small flowers? An experimental study of Brachyloma ericoides (Epacridaceae)

Abstract In many previous studies of the effects of introduced honeybees on Australian ecosystems, it has been assumed that floral morphology is a primary factor determining whether introduced honeybees will be effective in pollinating endemic plants. Although both honeybees and birds contacted stigmas and anthers of the small‐flowered Brachyloma ericoides (Epacridaceae), the exclusion of birds but not honeybees resulted in a significantly lower proportion of flowers producing capsules (12.3 ± 2 vs 21.0 ± 2%). This suggests that native birds contributed significantly to fruit set even though honeybees were much more frequent visitors to flowers (5–6 vs 0.7–2.5 times per day) and moved more frequently between plants (25 vs 12.2% of movements). Fruit set following exposure to birds and honeybees was very low compared with shrub species in general and may have been limited by the pre‐emptive removal of pollen by the 10% of honeybees that actively collected pollen.     

Reproductive biology of the Red List species Polemonium caeruleum (Polemoniaceae)

The majority of flowering plants, including many rare and threatened species, are pollinated by animals, but little is known of pollination and breeding systems of many endangered species. Polemonium caeruleum (Polemoniaceae) is a red‐listed species and is regarded as dichogamous, self‐compatible and bee pollinated. However, some studies show that it is visited by a vast assemblage of anthophilous insects from many taxonomic orders and that breeding systems vary greatly between closely related taxa of this genus. Over a period of 3 years we investigated breeding system, dichogamy, nectar secretion and composition, insect visitations and pollen loads in flowers of P. caeruleum in north‐eastern Poland to determine whether the reproductive biology of the plant explains its rarity. Contrary to published data, our study plants were self‐incompatible and showed a high degree of outcrossing. Our experimental work confirmed the occurrence of protandry in this species, revealed that nectar is sucrose‐dominant and proline‐rich and, for the first time for Polemoniaceae, that nectar secretion and nectar sugar concentration in flowers of P. caeruleum is female‐biased. Although flowers were visited by at least 39 species of insects from five taxonomic orders, overall the plant exhibited many characters associated with bee pollination, and analysis of insect performance showed that bumblebees and honeybees are the key pollinators; occasionally hoverflies and butterflies may also be involved. We conclude that, in terms of pollination system, P. caeruleum demonstrates high apparent generalization, but low realized generalization, and is a functional specialist, as most pollinators belong to a single functional group (guild). Its conservation status, at least in our study population, cannot be explained in terms of the biological properties of its breeding or pollination systems; rather, the present decline of the species is caused by habitat loss. However, if this process and bumblebee decline in Europe continue, P. caeruleum populations may diminish in numbers and density and, owing to the self‐incompatibility of the species, quickly become severely pollen‐limited, thereby accelerating further local extinctions. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 92–107.     

Differentiation Patterns of Reproductive Systems in the Genus Trillium

Abstract The differentiation patterns of reproductive systems, including breeding and pollination systems, and perennation strategies of the genus Trillium were critically studied. The pedicellate- and sessile-flowered species with different floral morphologies and ecological distributions exhibited distinct modes of reproductive systems in relation to their habitat conditions. The pedicellate-flowered species occurring in stable mesic temperate deciduous forests reproduce exclusively by seeds, while sessile-flowered species growing in the alluvial flood plain habitats depend to a large extent on vegetative reproduction to compensate for the insufficient sexual reproduction under ecologically unstable conditions, e.g., frequent disturbance due to flooding. These sessile-flowered species are characterized by low ovule numbers, subsequent low seed outputs and low fecundity levels. In contrast to the sessile-flowered species, the pedicellate-erect-flowered species showed very high seed setting rates of over 50%. Breeding experiments for four Japanese species suggest that, although they substantially possess the capability of both inbreeding and outbreeding, the high fecundity levels are maintained by predominant inbreeding system. Furthermore, resource levels in the stock organs of these species obviously determine fecundity levels. As a consequence the amount of reserved food in the rhizomes transferred to the next season is determined, which gurantees a continuous reproductive activity of a given individual plant.     

Reproductive biology and pollination of Utricularia reniformis A.St.‐Hil. (Lentibulariaceae)

Utricularia reniformis is an endemic Brazilian carnivorous plant, most common in high‐altitude grasslands. Knowledge of the reproductive biology of U. reniformis is essential for planning conservation strategies, but it is currently poorly understood. Thus, we studied the floral morphology, floral biology, breeding system and pollination of this species. U. reniformis produces and stores nectar in the flower spur, a classic feature of bee‐pollinated flowers, and we recorded Xylocopa sp. and Bombus sp. as pollinators. Moreover, although it is self‐compatible it is an obligate animal‐pollinated species, as the sensitive stigma avoids self‐pollination. However, in natural conditions reproductive success is low due to the rarity of visits from pollinators. We suggest that the low reproductive success caused by a deficit of pollinators may affect gene flow, causing loss of genetic diversity in U. reniformis.     

Differences in female reproductive success between female and hermaphrodite individuals in the subdioecious shrub Eurya japonica (Theaceae)

Subdioecy is thought to occupy a transitional position in the gynodioecy–dioecy pathway, explaining one of the evolutionary routes from hermaphroditism to dioecy. Quantifying any female reproductive advantage of females versus hermaphrodites is fundamental to examining the spectrum between subdioecy and dioecy; however, this is challenging, as multiple interacting factors, such as pollen limitation and resource availability, affect plant reproduction. We compared the female reproductive success of females and hermaphrodites via a field experiment in which we hand‐pollinated individuals of the subdioecious shrub Eurya japonica of similar size growing under similar light conditions. Effects of pollen limitation and seed quality were also evaluated through comparing the results of hand‐ and natural‐pollination treatments and performing additional laboratory and greenhouse experiments. Overall, females had higher fruit set and produced heavier fruit and more seeds than hermaphrodites, and these results were more pronounced for hand‐pollinated than for natural‐pollinated plants of both sexes. We also found that seeds naturally produced by females had a higher mean germination rate. These results indicate that females had a pronounced advantage in female reproductive success under conditions of no pollen limitation. The sexual difference in the degree of pollen limitation suggests a pollinator‐mediated interaction, whereas the higher female reproductive success of females even under natural conditions implies that E. japonica is a good model species for elucidating the later stages of the gynodioecy–dioecy pathway.     

Reproductive success and effective population size in woodrats (Neotoma macrotis)

Discrepancies between the census size and the genetically effective size of populations (N(e)) can be caused by a number of behavioural and demographic factors operating within populations. Specifically, strong skew in male reproductive success, as would be expected in a polygynous mating system, could cause a substantial decrease in N(e) relative to census size. Because the mating system of Neotoma macrotis had previously been described as one nearing harem polygyny, I examined the distribution of reproductive success and genetic variation within a population of this species. Combining genetic data and three years of field observations, I show that variance in reproductive success does not deviate from poisson expectations within either sex and variance in success is similar between the sexes. Furthermore, both males and females had multiple partners across litters in addition to some evidence of multiple paternity within litters. Despite a lack of strong skew in reproductive success, an estimate of N(e) based on a number of demographic parameters suggests that the ratio of N(e)/N in this population is 0.48. Although the ratio of N(e)/N suggests that the population is experiencing higher rates of genetic drift than would be expected based on census size alone, the population maintains high levels of genetic diversity. Estimates of neighbourhood size and patterns of recruitment to the study site suggest that immigration plays an important role in this population and may contribute to the maintenance of high levels of genetic diversity.     

Costs of mating competition limit male lifetime breeding success in polygynous mammals

Although differences in breeding lifespan are an important source of variation in male fitness, the factors affecting the breeding tenure of males have seldom been explored. Here, we use cross-species comparisons to investigate the correlates of breeding lifespan in male mammals. Our results show that male breeding lifespan depends on the extent of polygyny, which reflects the relative intensity of competition for access to females. Males have relatively short breeding tenure in species where individuals have the potential to monopolize mating with multiple females, and longer ones where individuals defend one female at a time. Male breeding tenure is also shorter in species in which females breed frequently than in those where females breed less frequently, suggesting that the costs of guarding females may contribute to limiting tenure length. As a consequence of these relationships, estimates of skew in male breeding success within seasons overestimate skew calculated across the lifetime and, in several polygynous species, variance in lifetime breeding success is not substantially higher in males than in females.