How seed traits predict floating times: a biophysical process model for hydrochorous seed transport behaviour in fluvial systems

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A non‐pollinating moth inflicts higher seed predation than two co‐pollinators in an obligate pollination mutualism

1. Mutualisms are relationships of mutual exploitation, in which interacting species receive a net benefit from their association. In obligate pollination mutualisms (OPMs), female pollinators move pollen between the flowers of a single plant species and oviposit eggs within the female flowers that they visit. 2. Competition between co‐occurring pollinator species is predicted to increase pollinator virulence, i.e. laying more eggs or consuming more seeds per fruit. Plants involved in OPMs frequently host various non‐pollinating seed parasites and parasitoids that may influence the outcome of the mutualism. Quantifying the prevalence of parasites and parasitoids and competition between pollinators is important for understanding the factors that influence OPM evolutionary stability. 3. This study investigated the pollination mutualism occurring between the leaf flower plant, Breynia oblongifolia, and its co‐pollinating Epicephala moths. A third moth, Herpystis, also occurs in B. oblongifolia fruits as a non‐pollinating seed parasite. 4. Breynia oblongifolia fruits were collected to quantify seed predation and compare seed predation costs between the three moth species. Results showed that the larvae of the two pollinator species consume similar numbers of seeds, and that adults deposit similar numbers of eggs per flower. As such, no evidence of increases in virulent behaviours was detected as a result of competition between co‐pollinators. 5. By contrast, the seed parasite Herpystis consumed more seeds than either pollinator species, and fruit crops with a high proportion of Herpystis had significantly lower net seed production. 6. This work adds to the growing understanding of the ecology and dynamics of plant–pollinator mutualisms.     

Sensitivity to Seed Germination Schedule by Scatter‐Hoarding Pére David's Rock Squirrels During Mast and Non‐Mast Years

Mast seeding by oaks (Quercus) and other seed species has fundamental impacts on the behavior of individual scatter‐hoarding animals as well as their population and community dynamics. Scatter‐hoarding squirrels are highly sensitive to acorn germination schedule and stop germination by removing the embryo of non‐dormant acorns to prevent losses of energy and nutrients by rapid germination. However, we know little about how this behavior is affected by mast seeding. We investigated foraging decisions made by free‐ranging Pére David's rock squirrel (Sciurotamias davidianus) with three species that produce non‐dormant seeds and two species that produce dormant seeds (Fagaceae) during mast and non‐mast years in Central China from 2007–2010. Consistent with the food perishability hypothesis that squirrels make decisions to minimize the perishability of their caches, non‐dormant seeds were hoarded at a lower rate, but had a higher probability of embryo removal than dormant seeds. Compared with non‐mast years, a lower proportion of seeds were harvested during mast years, but a higher proportion of the harvested seeds were hoarded. In addition, the probability of embryo removal in the hoarded seeds was higher during mast years. Squirrels also dispersed seeds at shorter distances during mast years. Moreover, the interactions between mast seeding and seed germination schedule had significant effects on squirrel foraging decisions, including embryo removal. This study is the first to demonstrate that squirrels show greater sensitivity to seed germination potential when seeds are abundant.     

Plant connectivity underlies plant‐pollinator‐exploiter distributions in Ficus petiolaris and associated pollinating and non‐pollinating fig wasps

Mutualism is ubiquitous in nature, and nursery pollination mutualisms provide a system well suited to quantifying the benefits and costs of symbiotic interactions. In nursery pollination mutualisms, pollinators reproduce within the inflorescence they pollinate, with benefits and costs being measured in the numbers of pollinator offspring and seeds produced. This type of mutualism is also typically exploited by seed‐consuming non‐pollinators that obtain resources from plants without providing pollination services. Theory predicts that the rate at which pollen‐bearing ‘foundresses’ visit a plant will strongly affect the plant's production of pollinator offspring, non‐pollinator offspring, and seeds. Spatially aggregated plants are predicted to have high rates of foundress visitation, increasing pollinator and seed production, and decreasing non‐pollinator production; very high foundress visitation may also decrease seed production indirectly through the production of pollinators. Working with a nursery mutualism comprised of the Sonoran Desert rock fig, Ficus petiolaris, and host‐specific pollinating and non‐pollinating fig wasps, we use linear models to evaluate four hypotheses linking species interactions to benefits and costs: 1) foundress density increases with host‐tree connectivity, 2) pollinator production increases with foundress density, and 3) non‐pollinator production and 4) seed production decrease with pollinator production. We also directly test how tree connectivity affects non‐pollinator production. We find strong support for our four hypotheses, and we conclude that tree connectivity is a key driver of foundress visitation, thereby strongly affecting spatial distributions in the F. petiolaris community. We also find that foundress visitation decreases at the northernmost edge of the F. petiolaris range. Finally, we find species‐specific effects of tree connectivity on non‐pollinators to be strongly correlated with previously estimated non‐pollinator dispersal abilities. We conclude that plant connectivity is highly important for predicting plant‐pollinator‐exploiter dynamics, and discuss the implications of our results for species coexistence and adaptation.     

Destination‐based seed dispersal homogenizes genetic structure of a tropical palm

As the dominant seed dispersal agents in many ecosystems, frugivorous animals profoundly impact gene movement and fine‐scale genetic structure of plants. Most frugivores engage in some form of destination‐based dispersal, in that they move seeds towards specific destinations, resulting in clumped distributions of seeds away from the source tree. Molecular analyses of dispersed seeds and seedlings suggest that destination‐based dispersal may often yield clusters of maternal genotypes and lead to pronounced local genetic structure. The long‐wattled umbrellabird Cephalopterus penduliger is a frugivorous bird whose lek mating system creates a species‐specific pattern of seed dispersal that can potentially be distinguished from background dispersal processes. We used this system to test how destination‐based dispersal by umbrellabirds into the lek affects gene movement and genetic structure of one of their preferred food sources Oenocarpus bataua, a canopy palm tree. Relative to background dispersal processes, umbrellabird mating behaviour yielded more diverse seed pools in leks that included on average five times more seed sources and a higher incidence of long‐distance dispersal events. This resulted in markedly lower fine‐scale spatial genetic structure among established seedlings in leks than background areas. These species‐specific impacts of destination‐based dispersal illustrate how detailed knowledge of disperser behaviour can elucidate the mechanistic link driving observed patterns of seed movement and genetic structure.     

Subcaste‐specific evolution of head size in the ant genus Pheidole

An organism's morphology is constrained by its evolutionary history and the need to meet a variety of potentially competing functions. The ant genus Pheidole is the most species‐rich ant genus and almost every species has a dimorphic worker caste (a few are trimorphic). This separation of workers into two developmentally distinct subcastes (smaller minors and larger majors with distinctively large heads) may partially release individuals from functional constraints on morphology, making Pheidole an ideal genus for addressing questions on the evolution of morphology in relation to ecological specialization. Major workers can perform a variety of tasks, although they are usually specialized for defence, as well as food retrieval and processing. Pheidole species vary in their diet, although many species gather seeds. The major workers mill the seeds using large jaws powered by mandible closer muscles that occupy a large proportion of the head cavity. In the present study, we examined the relationship between seed‐harvesting and morphology in Pheidole, hypothesizing that majors of seed‐harvesting species would have larger heads relative to non‐seed‐harvesters to accommodate the powerful mandibular muscles needed to mill seeds. By taking a phylogenetically controlled comparative approach, we found that majors of seed‐associated Pheidole did not have larger heads (width and length) than majors of non‐seed‐harvesting species. However, the head length of minors (and to a lesser extent head width) was smaller in seed‐harvesters. Additionally, we found the difference in head size between majors and minors was greater in seed‐harvesting species. These morphological differences in diet, however, were not related to changes in the rate of evolution in either seed‐harvesting or non‐seed‐harvesting lineages. These findings suggest that the morphologies of worker subcastes can evolve independently of each other, allowing colonies with polymorphic workers to specialize on new resources or tasks in ways not possible in monomorphic species.     

Endozoochory largely outweighs epizoochory in migrating passerines

Fruits and seeds are critical food sources for many European passerines during the autumn migration, which in turn contribute to disperse seeds either internally, i.e. after ingestion (endozoochory), or externally, when seeds adhere to the body surface (epizoochory). Despite the recognized importance of birds as seed dispersers, the vast majority of studies focused on endozoochory while the external transport of seeds is frequently invoked as being potentially important, but remains largely unexplored. This is particularly important during the post‐breeding migration of passerines, the most ubiquitous and diverse movement of potential seed carriers across Europe and into Africa, which coincides with the fruiting peak of many plant species (August–October). Our aim was to evaluate the role of migrating birds as potential long‐distance seed dispersers, and comparing the prevalence of epizoochory and endozoochory during post‐breeding migration. We sampled 926 wild birds in nine locations in Portugal, and retrieved 1833 seeds of 19 plant species dispersed internally and only three seeds externally attached to three birds (Serinus serinus, Locustella naevia and Turdus merula), showing an endozoochory prevalence 85 times higher than that of epizoochory. Migrating and non‐migrating passerines dispersed seeds equally. While two of the seeds transported externally had specific adaptations to epizoochory, namely spines (Torilis arvensis) and hooks (Galium aparine), the third is a large seed from a fleshy‐fruited plant, Frangula alnus (i.e. typical endozoochorous syndrome). These seeds were found on bird species with different diets, but similar behaviour (ground foragers) and in similar habitats (open agro‐ecosystems). Our results highlight the strong role of migrating passerines as potentially long‐distance seed dispersers and show that, at least in the autumn, the prevalence of epizoochory is several orders of magnitude lower than that of endozoochory.     

A graphical method for identifying the six types of non‐deep physiological dormancy in seeds

We present a new seed dormancy classification scheme for the non‐deep level of the class physiological dormancy (PD), which contains six types. Non‐deep PD is divided into two sublevels: one for seeds that exhibit a dormancy continuum (types 1, 2 and 3) and the other for those that do not exhibit a dormancy continuum (types 4, 5 and 6). Analysis of previous studies showed that different types of non‐deep PD also can be identified using a graphical method. Seeds with a dormancy (D) ? conditional dormancy (CD) ? non‐dormancy (ND) cycle have a low germination percentage in the early stages of CD, and during dormancy loss the germination capacity increases. However, seeds with a CD/ND (i.e. D→CD?ND) cycle germinate to a high percentage at a narrow range of temperatures in the early stages of CD. Cardinal temperatures for seeds with either a D/ND or a CD/ND cycle change during dormancy loss: the ceiling temperature increases in seeds with Type 1, the base temperature decreases in seeds with Type 2 and the base and ceiling temperatures decrease and increase, respectively, in seeds with Type 3. Criteria for distinguishing the six types of non‐deep PD and models of the temperature functions of seeds with types 1, 2 and 3 with both types of dormancy cycles are presented. The relevancy of our results to modelling the timing of weed seedling emergence is briefly discussed.     

Seed dispersal effectiveness by a large‐bodied invasive species in defaunated landscapes

Animal‐dispersed plants are increasingly reliant on effective seed dispersal provided by small‐bodied frugivores in defaunated habitats. In the Neotropical region, the non‐native wild pig (Sus scrofa) is expanding its distribution and we hypothesized that they can be a surrogate for seed dispersal services lost by defaunation. We performed a thorough analysis of their interaction patterns, interaction frequencies, seed viability, and characteristics of the seed shadows they produce. We found 15,087 intact seeds in 56% of the stomachs and 5,186 intact seeds in 90% of the scats analyzed, 95% of which were smaller than 10 mm in diameter. Wild pigs were the third most effective disperser among 21 extant frugivore species in a feeding trail experiment in terms of quantity of seeds removed. Gut retention time was 70 ± 23 hr, indicating wild pigs can promote long‐distance seed dispersal. Seed survival after seed handling and gut passage by wild pigs was positively related with seed size, but large seeds were spat out and only smaller seeds were defecated intact, for which we observed a positive or neutral effect on germination relative to manually de‐pulped seeds. Finally, deposition of seeds was four times more frequent in unsuitable than suitable sites for seedling recruitment and establishment. Seed dispersal effectiveness by wild pigs is high in terms of the quantity of seeds dispersed but variable in terms of the quality of the service provided. Our study highlights that negative and positive effects delivered by non‐native species should be examined in a case by case scenario. Abstract in Portuguese is available with online material.     

Is the pollen‐limited mistletoe Peraxilla tetrapetala (Loranthaceae) also seed limited?

Abstract Seed production in the endemic New Zealand mistletoe Peraxilla tetrapetala (Loranthaceae) has been shown to be consistently pollen limited, but to date there has been no test of whether the species is also seed limited. If it is not seed limited, then pollen limitation may have no effect on population size. We tested for seed limitation by sowing seeds onto host trees at high and low densities at two sites, and following survival for up to 69 months. Our rationale was that unless there is strong density‐dependent seedling mortality sufficient to negate increases in seed supply, the species would be seed limited. We simultaneously measured pollen limitation at both sites using pollen augmentation. Peraxilla tetrapetala was strongly pollen limited at both sites over 7 years at Ohau (mean Pollen Limitation Index 0.62), and 12 years at Craigieburn (mean PLI 0.44), before and during the seed‐limitation study. There was no significant overall negative effect of density on survival in sown P. tetrapetala seeds over 42–69 months. There was a significant positive effect: seeds were more likely to adhere to branches when sown at high densities. This initial advantage to high‐density seeds was gradually eroded by slightly (but non‐significantly) lower survival rates of adhered seeds at high density. By the end of the study there was no significant difference in overall survival in high and low density plantings. This means that P. tetrapetala was apparently both pollen limited and seed limited at both our sites. Hence, reduced densities of native bird pollinators caused by introduced mammalian carnivores are likely to reduce the density of adult mistletoes in the next generation. However, the generality of this result may be affected by the fact that mistletoes do not have a dense ‘seedling shadow’ under the parent mistletoe.     

Nutrient transport within and between habitats through seed dispersal processes by woolly monkeys in north‐western Amazonia

The contribution of vertebrate animals to nutrient cycling has proven to be important in various ecosystems. However, the role of large bodied primates in nutrient transport in neotropical forests is not well documented. Here, we assess the role of a population of woolly monkeys (Lagothrix lagothricha lugens) as vectors of nutrient movement through seed dispersal. We estimated total seed biomass transported by the population within and between two habitats (terra firme and flooded forests) at Tinigua Park, Colombia, and quantified potassium (K), phosphorus (P) and nitrogen (N) content in seeds of 20 plant species from both forests. Overall, the population transported an estimated minimum of 11.5 (±1.2 SD) g of potassium, 13.2 (±0.7) g of phosphorus and 34.3 (±0.1) g nitrogen, within 22.4 (±2.0) kg of seeds ha^?1 y^?1. Approximately 84% of all nutrients were deposited in the terra firme forest mostly through recycling processes, and also through translocation from the flooded forest. This type of translocation represents an important and high‐quality route of transport since abiotic mechanisms do not usually move nutrients upwards, and since chemical tests show that seeds from flooded forests have comparatively higher nutrient contents. The overall contribution to nutrient movement by the population of woolly monkeys is significant because of the large amount of biomass transported, and the high phosphorus content of seeds. As a result, the phosphorus input generated by these monkeys is of the same order of magnitude as other abiotic mechanisms of nutrient transport such as atmospheric deposition and some weathering processes. Our results suggest that via seed dispersal processes, woolly monkey populations can contribute to nutrient movement in tropical forests, and may act as important nutrient input vectors in terra firme forests. Am. J. Primatol. 72:992–1003, 2010. © 2010 Wiley‐Liss, Inc.     

Mixed pollen load and late‐acting self‐incompatibility flexibility in Adenocalymma peregrinum (Miers) L.G. Lohmann (Bignonieae: Bignoniaceae)

• Mixed cross and self‐pollen load on the stigma (mixed pollination) of species with late‐acting self‐incompatibility system (LSI) can lead to self‐fertilized seed production. This “cryptic self‐fertility” may allow selfed seedling development in species otherwise largely self‐sterile. Our aims were to check if mixed pollinations would lead to fruit set in LSI Adenocalymma peregrinum, and test for evidence of early‐acting inbreeding depression in putative selfed seeds from mixed pollinations.
• Experimental pollinations were carried out in a natural population. Fruit and seed set from self‐, cross and mixed pollinations were analysed. Further germination tests were carried out for the seeds obtained from treatments.
• Our results confirm self‐incompatibility, and fruit set from cross‐pollinations was three‐fold that from mixed pollinations. This low fruit set in mixed pollinations is most likely due to a greater number of self‐ than cross‐fertilized ovules, which promotes LSI action and pistil abortion. Likewise, higher percentage of empty seeds in surviving fruits from mixed pollinations compared with cross‐pollinations is probably due to ovule discounting caused by self‐fertilization. Moreover, germinability of seeds with developed embryos was lower in fruits from mixed than from cross‐pollinations, and the non‐viable seeds from mixed pollinations showed one‐third of the mass of those from cross‐pollinations.
• The great number of empty seeds, lower germinability, lower mass of non‐viable seeds, and higher variation in seed mass distribution in mixed pollinations, strongly suggests early‐acing inbreeding depression in putative selfed seeds. In this sense, LSI and inbreeding depression acting together probably constrain self‐fertilized seedling establishment in A. peregrinum.

     

Movement Patterns of a Native and Non‐native Frugivore in Hawaii and Implications for Seed Dispersal

Avian frugivores historically played important roles as seed dispersers across the Hawaiian Islands, but presently, the ‘ōma‘o (Myadestes obscurus) is the only extant native frugivore in the wild on the Island of Hawaii. During recent decades, the introduced generalist Japanese White‐eye (Zosterops japonicus) has become the most common bird in Hawaii. The movements of avian frugivores largely dictate how far seeds get dispersed and into what kinds of microhabitats. This study compares the movement patterns and diet of the ‘ōma‘o to the Japanese White‐eye to understand how a native differs from a non‐native frugivore in the type and distances of seeds dispersed. Radiotelemetry was conducted on nine ‘ōma‘o and nine Japanese White‐eyes in a system of natural forest fragments (kīpuka) created by lava flows. Japanese White‐eyes disperse seeds approximately twice as far as ‘ōma‘o; during the time of gut passage, ‘ōma‘o move a mean distance of 98.1 m, and Japanese White‐eyes move 170.1–194.8 m. However, the ‘ōma‘o disperses the seeds of at least seven different native fruit species compared with two dispersed by Japanese White‐eyes. Japanese White‐eyes were found to disperse seeds smaller than 1.5 mm, whereas the ‘ōma‘o dispersed seeds up to 6 mm in diameter. Despite their ecological differences, both birds distribute certain seeds within and among kīpuka and likely facilitate primary succession of fruiting plants in the young lava matrix. However, this study suggests that if the ‘ōma‘o were extirpated, a smaller‐bodied generalist cannot entirely substitute for the ecological role played by the native frugivore.     

Costs and benefits of non‐random seed release for long‐distance dispersal in wind‐dispersed plant species

The dispersal ability of plants is a major factor driving ecological responses to global change. In wind‐dispersed plant species, non‐random seed release in relation to wind speeds has been identified as a major determinant of dispersal distances. However, little information is available about the costs and benefits of non‐random abscission and the consequences of timing for dispersal distances. We asked: 1) to what extent is non‐random abscission able to promote long‐distance dispersal and what is the effect of potentially increased pre‐dispersal risk costs? 2) Which meteorological factors and respective timescales are important for maximizing dispersal? These questions were addressed by combining a mechanistic modelling approach and field data collection for herbaceous wind‐dispersed species. Model optimization with a dynamic dispersal approach using measured hourly wind speed showed that plants can increase long‐distance dispersal by developing a hard wind speed threshold below which no seeds are released. At the same time, increased risk costs limit the possibilities for dispersal distance gain and reduce the optimum level of the wind speed threshold, in our case (under representative Dutch meteorological conditions) to a threshold of 5–6 m s^–1. The frequency and predictability (auto‐correlation in time) of pre‐dispersal seed‐loss had a major impact on optimal non‐random abscission functions and resulting dispersal distances. We observed a similar, but more gradual, bias towards higher wind speeds in six out of seven wind‐dispersed species under natural conditions. This confirmed that non‐random abscission exists in many species and that, under local Dutch meteorological conditions, abscission was biased towards winds exceeding 5–6 m s^–1. We conclude that timing of seed release can vastly enhance dispersal distances in wind‐dispersed species, but increased risk costs may greatly limit the benefits of selecting wind conditions for long‐distance dispersal, leading to moderate seed abscission thresholds, depending on local meteorological conditions and disturbances.     

Paddock‐scale patterns of seed production and dispersal in the invasive shrub Acacia nilotica (Mimosaceae) in northern Australian rangelands

Paddock‐scale Acacia nilotica L. Willd. ex Del. (prickly acacia) spatial distribution, seed production and dispersal patterns were investigated in the Astrebla (Mitchell) grasslands of northern Australia as a step toward predicting future patterns of invasion. A number of hypotheses were tested based on what we know of this species in both Australia and regions where it is native, for example South Africa. It was hypothesized that most A. nilotica seeds would be produced by trees in riparian habitats with access to permanent water. In addition, we predicted that seeds would be dispersed throughout the Astrebla grassland landscape by cattle, following observations that cattle readily ingest and pass seeds and that cattle have access to all areas within paddock boundary fences. Tree density, seed production and seed dispersal by cattle were measured along a series of transects from permanent watering points to paddock boundary fences. Trees associated with permanent watering points produced more seeds per unit area and occurred at higher density than their non‐riparian counterparts. The importance of riparian trees decreased in years with high rainfall and in paddocks with only small areas of riparian habitat. Cattle spread dung and seeds throughout paddocks, with peaks of deposition adjacent to permanent watering points. These results suggest that invasion patterns are likely to be uneven across the landscape and may be reactive to climate. High seedling recruitment and possible thicket formation is expected adjacent to permanent watering points and wherever cattle congregate. Patterns of recruitment in non‐riparian areas are likely to be relatively sparse. The importance of post‐dispersal factors in determining recruitment patterns is discussed.     

A biotechnology‐based male‐sterility system for hybrid seed production in tomato

Incorporating male sterility into hybrid seed production reduces its cost and ensures high varietal purity. Despite these advantages, male‐sterile lines have not been widely used to produce tomato (Solanum lycopersicum) hybrid seeds. We describe the development of a biotechnology‐based breeding platform that utilized genic male sterility to produce hybrid seeds. In this platform, we generated a novel male‐sterile tomato line by clustered regularly interspaced short palindromic repeat (CRISPR)/CRISPR‐associated protein 9 (Cas9)‐mediated mutagenesis of a stamen‐specific gene SlSTR1 and devised a transgenic maintainer by transforming male‐sterile plants with a fertility‐restoration gene linked to a seedling‐colour gene. Offspring of crosses between a hemizygous maintainer and the homozygous male‐sterile plant segregated into 50% non‐transgenic male‐sterile plants and 50% male‐fertile maintainer plants, which could be easily distinguished by seedling colour. This system has great practical potential for hybrid seed breeding and production as it overcomes the problems intrinsic to other male‐sterility systems and can be easily adapted for a range of tomato cultivars and diverse vegetable crops.     

Experimental determination of seed emergence and carry‐over in the soil seed bank of the herbaceous perennial,Trachymene incisa (Apiaceae)

Abstract Many populations of herbaceous perennial plants contain seeds stored in a soil seed bank. The contribution of seeds to population persistence is an important parameter in population models but germination rates of known‐age seeds are difficult to obtain because individual seeds cannot easily be followed. Although Trachymene incisa Rudge plants produce copious seeds that are dispersed into the soil, the existence of a seed bank has not been confirmed. To quantify the potential for a seed bank fresh seeds of T. incisa were sown into experimental seed banks in the eucalypt‐dominated Agnes Banks Woodland in western Sydney, NSW. A recent fire provided the opportunity to compare germination in the burnt and unburnt vegetation. Density of seed sowing and time of maturation/dispersal of seeds were manipulated in 75 seed cages. Emergence of seeds after 5 months was significantly higher for the earliest planting date but after 1 year, germination of seeds planted in the later weeks increased, and the final germination for all weeks was 28%. Density of sowing and the recent fire did not affect emergence. A second experiment planted over a broader time span (9 weeks instead of 3 weeks) confirmed the effect of planting date but also found significant spatial variation on a scale of tens of metres. Laboratory germination rates of over 70% confirmed that the seeds were viable and non‐dormant when sown in the field cages. The carry‐over of non‐germinated seed in the soil seed bank is estimated to be about 70% after 2 years, implying that a cohort of seeds would not be depleted through germination alone for up to 40 years. The potential for a long‐lived seed bank in this species is interesting because the plants are also capable of resprouting from their rootstock after fire, giving them characteristics of both resprouters and seeders.