Dung beetles as secondary seed dispersers in a temperate grassland

The two-phase dispersal event in which dung beetles move seeds after endozoochory is often assumed to be advantageous for plant regeneration. Because seeds are expected to end up in favourable and safe germination sites, it is considered as an example of directed dispersal. However, literature so far is restricted to tropical rain forest ecosystems, while data for temperate regions are lacking. In this study, the effect of dung beetles on seedling establishment of endozoochorically dispersed seeds is evaluated for a temperate grassland ecosystem. We performed a field experiment in which cages excluded dung beetles from horse and cattle dung samples with mixed-in grass seeds. Seed germination from these samples was significantly higher than that from samples which were accessible to dung beetles. This indicates that the effect of dung beetles on short-term seedling establishment was negative, which contrasts with the patterns found for large-seeded species used in tropical studies. This is most likely attributed to the lack of roller species and the larger depth at which tunneling Geotrupes species bury seeds.     

Establishment of pasture species on cattle dung: the role of endozoochorous seeds

Abstract. The process of colonization of cattle dung patches and its relation to endozoochorous dispersal is analysed in a Mediterranean pasture. Dung pats dropped in spring and winter were marked, and the vegetation developing on them during the first two years of colonization was followed. The influence of endozoochorous seeds on dung colonization is assessed through the study of: (1) the vegetation on spring and winter dung pats and the seed contents of both, which is abundant in the former and negligible in the latter; and (2) the vegetation on spring dung pats compared with that of the surrounding pasture. The origin of plants growing on dung (either seeds in the manure or in the soil seed bank) was traced by studying dung colonization under controlled conditions in pots with manure put on seed-free sand, and on pasture turf. The results indicate that endozoochorous seeds germinate in manure; they are the main source of recovery in gaps generated by dung pats. The micro-succession involved is independent of the type of pasture. A small-scale spatial pattern results in which gaps of old dung are dominated by endozoochorous species. Thus, dung patches enhance the similarity between grazed communities, and the variation within communities.     

Why are small seeds dispersed through animal guts: large numbers or seed size per se?

Long-distance dispersal of seeds is an important process in metapopulation dynamics and in plant migrations, but at the same time extremely difficult to observe or quantify directly. If seed dispersal ability were related to attributes of seeds or motherplants, long-distance seed dispersal would be predictable by indirect approximation using easy-to-measure traits. Seed size has been suggested to be such a key trait in seed dispersal ability. However, having smaller seeds also implies having more numerous seeds per plant individual (given equal reproductive effort), and consequently increases the probability of seeds being ingested accidentally. The question is whether small-seeded species are more abundant in herbivore dung because smaller seed size increases survival rate during gut passage or because they are produced (and ingested) in greater numbers than larger seeds. We investigated endozoochorous seed dispersal via cattle grazing a meadow, and related seed abundance in dung samples to seed attributes. We found that seeds were ingested and passed through the bovine intestinal tract in proportion to the numbers produced per unit area in the grazed vegetation. In contrast, no relationship could be found between endozoochorous dispersal potential (measured as abundance of seeds in dung samples corrected for seed output in the grazed vegetation) and seed attributes such as seed mass, seed shape (roundness), and thickness of the seed coat. This finding underlines the importance of seed number in plant dispersal ability. In addition, it shows that grazing mammals may constitute an important dispersal vector for many plant species conventionally classified as 'unspecialised'.     

Endozoochorous seed dispersal by free-ranging herbivores in an abandoned landscape

Seed dispersal is a crucial process for the dynamics and maintenance of plant populations. Free-ranging animals are effective dispersal vectors because they can move between similar habitats and transport seeds into favourable environments. Dung samples from two species of common free-ranging mammals—deer and wild boar—were used to study endozoochorous dispersal of seeds in a military training area in western Bohemia. The area was abandoned after WWII, and the military training area was established in 1953. The vegetation consists of shrublands and dry grasslands. Data on the local species pool of grassland herbs and forbs were collected to compare the characteristics of dispersed versus non-dispersed plants. Deer and wild boar dispersed 84 plant species; however, species composition of seedlings emerging from dung samples showed significant differences between dispersal vectors and notable change across the growing season. 80% of all seedlings extracted from the dung samples belonged to stinging nettle, Urtica dioica. From trait analyses, seeds of endozoochorous plants had a higher longevity index in the soil seed bank than non-endozoochorous plants and more often possessed a mucilaginous surface. Our results show that deer and boar are successful, though not substitutable dispersers.     

Seed bank build-up in small disturbances in a Mediterranean pasture: the contribution of endozoochorous dispersal by rabbits

The importance of endozoochorous dispersal in seed bank build-up is analyzed in artificial 40 x 40 cm disturbances in a Mediterranean pasture The top 10 cm of soil in the disturbances was replaced by sterile soil, and the excrement laid m half of them was removed fortnightly The seed banks of the disturbances subjected to the two treatments was quantified by germination of soil samples (n = 44) gathered at the start of the following growth season Rabbit droppings were collected fortnightly to quantify their seed content and the pasture seed production was estimated Seed bank regeneration takes place early in the disturbances The main source of qualitative variation in seed banks is their position on the slope, while endozoochory lacks a characteristic qualitative effect However, endozoochorous dispersal is responsible for a significant increase in the number of germinable seeds, the richness and diversity of the seed bank samples with respect to those lacking it The importance of endozoochorous dispersal by rabbits in grassland dynamics is suggested to be its effect on the colonization of small disturbances, where species dispersed with dung are found to be favoured This aids the maintenance of grazing-tolerant communities     

Impact of the spatial uncertainty of seed dispersal on tree colonization dynamics in a temperate forest

An aggregated distribution of dispersed seeds may influence the colonization process in tree communities via inflated spatial uncertainty. To evaluate this possibility, we studied 10 tree species in a temperate forest: one primarily barochorous, six anemochorous and two endozoochorous species. A statistical model was developed by combining an empirical seed dispersal kernel with a gamma distribution of seedfall density, with parameters that vary with distance. In the probability density, the fitted models showed that seeds of Fagaceae (primarily barochorous) and Betulaceae (anemochorous) were disseminated locally (i.e. within 60 m of a mother tree), whereas seeds of Acer (anemochorous) and endozoochorous species were transported farther. Greater fecundity compensated for the lower probability of seed dispersal over long distances for some species. Spatial uncertainty in seedfall density was much greater within 60 m of a mother tree than farther away, irrespective of dispersal mode, suggesting that seed dispersal is particularly aggregated in the vicinity of mother trees. Simulation results suggested that such seed dispersal patterns could lead to sites in the vicinity of a tree being occupied by other species that disperse seeds from far away. We speculate that this process could promote coexistence by making the colonization rates of the species more similar on average and equalizing species fitness in this temperate forest community.     

Seed dispersal in the mycoheterotrophic orchid Yoania japonica: Further evidence for endozoochory by camel crickets

• Although orthopterans are rarely considered to be effective seed dispersal agents, the large flightless crickets known as ‘weta’ have been suggested to function as ecological replacements for small mammals in New Zealand, where such mammals are absent. In addition, a recent study reported that camel crickets mediate seed dispersal of several heterotrophic plants, including Yoania amagiensis in Japan.
• I investigated the seed dispersal mechanism of Yoania japonica because the fruit morphology is similar to Y. amagiensis. Specifically, I aimed to determine whether Y. japonica fruits are consumed by camel crickets and, if so, whether the seeds defecated by camel crickets remains intact, by checking seed viability with TTC staining, and whether germination rate is different between seeds collected directly from fruits and defecated seeds by comparing in situ seed germinability.
• The present study provides evidence that camel crickets function as seed dispersal agents of Y. japonica. Camel crickets were important consumers of Y. japonica fruits, and a substantial portion of the consumed seeds remained viable after passing through the digestive tract. In situ seed germination experiments revealed that the seeds defecated by camel crickets actually germinated in the field. In addition, the germination rate of defecated seeds was even higher than that of intact seeds, although the difference was not significant.
• Taken together with recent reports of insect‐mediated endozoochory, such a seed dispersal system may be common in plants with fleshy indehiscent fruits and small seeds, even in locations where other seed dispersal agents are present.

     

Endozoochorous seed dispersal by cattle and horse in a spatially heterogeneous landscape

Seed dispersal has become an important issue in plant ecology and restoration management. In this paper we examined dung germinating seed content and seed deposition patterns of horses (Shetland and Konik breeds) and Scottish Highland cattle grazing two coastal dune nature reserves. Two times 2.5 l of fresh dung from each type of herbivore were collected during seven sessions in the main fruiting season. Dung samples were placed under greenhouse conditions after drying and cooling. Animal defecation patterns were derived from a study of herbivore activities during 6 h observation sessions 8 times a month. One hundred and seventeen plant species i.e. 27% of all species occurring in the study area, were recorded as seedlings emerging from the dung samples. The most abundantly and frequently recorded plant species were Urtica dioica, Juncus spp. and different species of Poaceae and Caryophyllaceae. In general seedling density is high (1158 seedlings/dung sample). Seedling density and species richness were further analysed in order to detect temporal variability and possible animal and site related characteristics. Dung deposition patterns reflect a non-random use of habitats and hence a non-random seed deposition among habitats. Calculated seed deposition per square meter ranged from a few (<10 germinating seeds) to more than 100 in the most frequently selected habitats. From the herbivores’ selective habitat use and their estimated mean retention time we can further assume their ability for inter-habitat endozoochorous seed dispersal. This characteristic of large herbivores is further discussed in the light of nature management and restoration.Indra Lamoot is an aspirant of FWO-Flanders.     

Diaspore dispersal season influences root growth and seed traits,but not germination responses,of sympatric Piper species in a seasonal forest

• Seeds may differ in terms of dormancy, longevity, sensitivity to desiccation and dry mass, according to the timing (dry season/rainy season) of diaspore dispersal. In addition, seasonal variations in temperature and water availability can act as signals of the season during seed development, influencing germination responses and root growth. We evaluated the effects of temperature variations and water availability on germination parameters, root growth and seed traits of four coexisting Piper species in seasonal vegetation that differed in diaspore dispersal timing.
• Eight temperature treatments (15, 20, 23, 25, 28, 30, 35 °C, and alternate 30 °C–20 °C) and four induced water potentials (0, −0.3, −0.6 and −1.2 MPa) were used. The parameters germination onset, germination percentage (G%), mean germination time (MGT), root elongation, seed longevity during ex situ storage and dry mass of seeds were evaluated.
• Germination responses observed were independent of the diaspore dispersal timing, such as variations in germination onset, G% and MGT, both in temperature and water availability treatments. In contrast, root elongation, longevity and dry mass of seeds varied according to the time of diaspore dispersal.
• Our results corroborate the hypothesis that the timing of diaspore dispersal is an important factor in controlling the initial development of seedlings in seasonal vegetation, but not in germination responses. The predominance of negative effects of temperature increases and water deficit on root growth shows that the initial stages of plant development can be strongly impacted by these environmental factors.

     

Livestock grazing ingestion suppressed the dominant species population (Stipa aliena) germination: a laboratory experiment

Grazing has an important effect on population dynamics in grassland vegetation. This paper aims at providing information on the impact of livestock ingestion on seed germination in a Tibetan meadow. We conducted a laboratory germination experiment in which Stipa aliena seeds were treated by yak and Tibetan sheep rumen fluids and yak dung extract with undiluted, 1‐fold and 10‐fold diluted solutions. The results showed that seed germinability was restrained in all these treatments. In the control treatment, 66.2% of the seeds germinated, while there was almost no S. aliena germination in the undiluted and 1‐fold diluted solutions of both yak and Tibetan sheep rumen fluids. Yak dung extract had relatively less effect on seed germination. Additionally, the primary effect of rumen fluid seems to be that it kills part of the seeds rather than delaying germination. Seed death may be due to the acidity of the fluid or/and due to the effect of microorganisms in the fluid. The negative effects of livestock digestive juices on S. aliena germination may constitute a mechanism explaining the population dynamics in alpine meadows of the Qinghai‐Tibetan Plateau.     

Seed traits and bird species influence the dispersal parameters of wetland plants

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Cattle dung and the fate of Biserrula pelecinus L. (Leguminosae) in a Mediterranean pasture: seed dispersal, germination and recruitment

Seed dispersal of Biserrula pelecinus via cattle dung in a semiarid pasture is analysed by quantification of the viable seed content of cattle dung, assessment of the percentage of readily germinable seeds in growth chamber of seeds collected from the plant and from dung, monitoring of the on-field frequency of the species in dung pats and in surrounding (control) pasture plots the four springs after dung deposition, and quantification of the pasture area occupied by dung pats. Large amounts of seeds of the species are dispersed by cattle (up to 4 seeds g^-1 of dry manure in June 1991), and the germinability of the seeds increases significantly following their passage through the cattle gut (from 2.5% to 9.5%). In the first three springs after deposition, the frequency of adult plant of the species on dung almost tripled that found in the surrounding pasture. The effect on the pasture is scale-dependent and varies considerably between zones depending on the area covered by dung. Thus, presence on dung may explain an average of 8% of the frequency of the species in the pasture at a 10 times 10 cm scale, and up to 20% in some areas. The results are the first evidence of a herbaceous species being greatly favoured in a plant community by its dispersal through herbivore dung, a process that may have profound implications for the interpretation of plant-herbivore relations and on the evolution of plant traits.     

Does seed ingestion by bats enhance germination? A new meta‐analysis 15 years later

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Scale-dependent diversity effects of seed dispersal by a wild herbivore in fragmented grasslands

Dispersal limitation between habitat fragments is a known driver of landscape-scale biodiversity loss. In Europe, agricultural intensification during the twentieth century resulted in losses of both grassland habitat and traditional grassland seed dispersal vectors such as livestock. During the same period, populations of large wild herbivores have increased in the landscape. Usually studied in woodland ecosystems, these animals are found to disperse seeds from grasslands and other open habitats. We studied endozoochorous seed dispersal by roe deer (Capreolus capreolus) in fragmented grasslands and grassland remnants, comparing dispersed subcommunities of plant species to those in the established vegetation and the seed bank. A total of 652 seedlings of 67 species emerged from 219 samples of roe deer dung. This included many grassland species, and several local grassland specialists. Dispersal had potentially different effects on diversity at different spatial scales. Almost all sites received seeds of species not observed in the vegetation or seed bank at that site, suggesting that local diversity might not be dispersal limited. This pattern was less evident at the landscape scale, where fewer new species were introduced. Nonetheless, long-distance dispersal by large wild herbivores might still provide connectivity between fragmented habitats within a landscape in the areas in which they are active. Finally, as only a subset of the available species were found to disperse in space as well as time, the danger of future biodiversity loss might still exist in many isolated grassland habitats.     

A global review of weeds that can germinate from horse dung

To assess the diversity of weed seeds dispersed via horse dung, we reviewed 15 studies on seed germination from horse dung – six from Europe, four from North America, three from Australia and one study each from Africa and Central America. Seed from 249 species from 43 families have been identified germinating from horse dung. Almost two‐thirds of the species were forbs and 33% graminoids, with over half being perennials and 32% annuals. Nearly every species (totalling 99% of those reviewed) is considered a weed somewhere, with 47% recorded as invasive and 19% international environmental weeds. Of the 2739 non‐native plants that are naturalized in Australia, 156 have been shown to germinate from horse dung. This includes 16 of the 429 listed noxious weeds in Australia and two weeds of national significance. Seed from 105 of the 1596 invasive/noxious plant species in North America have also been identified germinating from horse dung. Seed traits including seed size, length, width and mass affect dispersal via horse dung. Habitat disturbance from trampling facilitates germination of seedlings from dung in both natural and experimental studies. Some studies found that plants germinating from dung reach maturity and flower, while others found plants did not survive due to unfavourable growing conditions in the field. The diversity of species with seed that can germinate from horse dung highlights the potential of horses to disperse a range of seed over long distances. Whether such dispersal is beneficial or harmful depends on the plant and the context in which it germinates. To maintain the conservation value of protected areas, it is important to understand and manage the different potential weed dispersal vectors, including horses.     

The effect of seeds of exotic species transported via horse dung on vegetation along trail corridors

It has been suggested that exotic species will colonize within forests more frequently by the continual introduction of seeds through horse dung deposited along trails. Whether or not these exotic species have the ability to spread into and establish in the forest interior has been disputed. To address this, horse dung and soil samples were collected from trails during Autumn 1994 and Summer 1995 from three areas in southern Illinois, USA open to recreational horse travel. In addition, deer dung samples were collected from each of the study areas. Vegetation data were collected from each of the trail systems as well as from a trail along which horse travel was prohibited. The density of vascular plants in 0.25 m² quadrats placed at varying distances from the trail center to 5 m into the forest interior were recorded. Finally, dung samples were placed in situ along horse trails at one site to examine seedling germination in natural conditions. While 23 exotic species germinated from samples of horse dung placed out in a greenhouse, only one of these exotic species was also found in trail plots (Kummerowia striata). Similarly, while there were empirically more exotic species found along the trails allowing horse travel than there were on the trail lacking horse travel, the relative importance of those species was negligible along both trails. These results suggest that the emigration of exotic species via horse dung does not pose an immediate threat to the plant communities adjacent to trails in these forest systems. Nevertheless, the large number of exotic species in horse dung reflects the constant threat to any system from these species. Care must be taken, when allowing horseback use in areas, to anticipate invasion by exotic species from horse dung     

Restoration of wooded meadows ‐ a comparative analysis along a chronosequence on Öland (Sweden)

Abstract. Wooded meadows on the Baltic Island of Öland result from traditional agricultural management over centuries which has led to a species‐rich vegetation with high species diversity. Today, nearly all of these meadows have been abandoned and became rapidly overgrown by deciduous shrub and tree species forming a closed canopy which resulted in a rapid and strong decrease in species numbers of the herb layer. Recent efforts aim to restore overgrown wooded meadows by cutting single shrubs and trees to open the canopy. However, the effects of abandonment as well as of any restoration management in wooded meadows have rarely been documented until now. Mechanisms driving succession after restoration such as the dispersal potential of the respective species over time and space have not been analysed yet. Therefore, a chronosequence was studied which included a traditionally managed wooded meadow, an overgrown meadow which has been abandoned for more than 100 yr and a meadow which was restored 36 yr ago by cutting and is now grazed. We analysed the soil seed bank of the 3 meadows in comparison with the established vegetation and endozoochorous seed dispersal by cattle and sheep. After abandonment 87% of the typical grassland species vanished from the established vegetation and were replaced by species characteristic of woodland and disturbed grassland communities. The mean number of species decreased from 52 species per plot (4 m²) to 18 species. Mean species number and number of seeds in the seed bank declined significantly from the traditionally managed to the overgrown meadow. Most of the grassland species were assigned to a transient seed bank type while only 1/3 could be classified as having a short‐term persistent seed bank. Thus, restoration of wooded meadows cannot rely on the soil seed bank. Endozoochorous seed dispersal by cattle and sheep was shown for 15% of the species with seed densities per 100 g air dried dung of 737 and 767, respectively. Movement of animals between ancient and restored wooded meadows is recommended since many of the species only occurred in low densities and therefore, will probably not be found in the dung samples.     

Heteromorphic seeds of wheat wild relatives show germination niche differentiation

• Crop wild relatives are fundamental genetic resources for crop improvement. Wheat wild relatives often produce heteromorphic seeds that differ in morphological and physiological traits. Several Aegilops and Triticum species possess, within the same spikelet, a dimorphic seed pair, with one seed being larger than the other. A comprehensive analysis is needed to understand which traits are involved in seed dimorphism and if these aspects of variation in dimorphic pairs are functionally related.
• To this end, dispersal units of Triticum urartu and five Aegilops species were X‐rayed and the different seed morphs weighed. Germination tests were carried out on seeds, both dehulled and left in their dispersal units. Controlled ageing tests were performed to detect differences in seed longevity among seed morphs, and the antioxidant profile was assessed in terms of antioxidant compounds equipment and expression of selected antioxidant genes. We used PCA to group seed morphs sharing similar patterns of germination traits, longevity estimates and antioxidant profile.
• Different seed morphs differed significantly in terms of mass, final germination, germination timing, longevity estimates and antioxidant profile in most of the tested species. Small seeds germinated slower, had lower germination when left in their dispersal units, a higher antioxidant potential and were longer‐lived than large seeds. The antioxidant gene expression varied between morphs, with different patterns across species but not clearly reflecting the phenotypic observations.
• The results highlight different trait trade‐offs in dimorphic seeds of Aegilops and T. urartu, affecting their germination phenology and longevity, thereby resulting in recruitment niche differentiation.

     

Seed germination in temperate rain forest species of southern Chile: chilling and gap-dependency germination

Canopy gap and chilling requirements for seed germination were assessed in 61 and 44 species, respectively, in the temperate rain forest of southern Chile. Germination assays within canopy gap and understorey were carried out under natural conditions. Germination tests in cold stratified and non-stratified seeds were performed under laboratory conditions. Seeds were collected of common trees, shrubs, vines and perennial herbs of forests in southern Chile. Final percent germination was significantly enhanced under canopy gap conditions in 19 species, and significantly reduced in 11 species. Germination proved indifferent under gap vs. understorey conditions in half the species tested. Cold stratification affected germination in 11 species: significantly increasing final germination of four species, and significantly decreasing final percent germination of seven species. A Principal Components Analysis (PCA) was applied with the purpose of identifying groups of species with similar germination strategies. Four attributes were included in the PCA: final germination percentage in canopy gap, germination rate in the laboratory, and gap-and chilling-dependency indices (EGAP and STRAT, respectively). The first axis separated species mainly on EGAP variation while the second axis separated them mainly according to STRAT variation; the two axes together explaining 73% of the among-species variation. A small group of trees and vines germinating best in the understorey and neutral to chilling could clearly be distinguished from the remaining species analyzed. Multifactorial ANOVAs were used for evaluating the combined effect of successional status, seed mass, dispersal period, life form, phylogenetic categories, and dispersal syndrome on EGAP and STRAT variation. The EGAP value of secondary successional species proved significantly greater than that of primary successional species and species with endozoochorous seed dispersal were significantly less dependent on chilling (according to STRAT values) compared to species with mainly wind-dispersed seeds. The possible implications of these results for seedling establishment are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of bird ingestion on seed germination of four woody species of the temperate rainforest of Chiloé island, Chile

We study the effect of ingestion by birds on seed germination and theconsequences of absence of dispersal, with the persistency of the seedsinside the fruit. We collected seeds of four woody species ofthe temperate rainforest of Chiloé: Gaultheriamucronata, Luma apiculata, Myrteolanummularia, and Myrceugenia planipes. The seedstested had the following origins: 1) Ingested seeds: seeds collected fromthe feces of birds, 2) Extracted seeds: seeds obtained directly from thefruits, and 3) Intact fruits: fruits collected directly from the plants.Germination of Myrceugenia planipes under greenhouseconditions, Luma apiculata, and Myrteolanummularia under laboratory conditions, and Gaultheriamucronata under both conditions was analyzed. We found that the seedsreach their maximum germination between 15–20 days after sowing, withthe exception of those of G. mucronata sown in the greenhouse,which showed a low germination rate. In the greenhouse assay, seeds ofG. mucronata ingested by birds, seeds extracted manuallyfrom the fruits, and seeds inside the fruits did not show significant differencesin their germination percentages. In the laboratory assays, the seeds ofG. mucronata and M. nummulariaingested by birds and the seeds extracted manually from the fruits also did not show anysignificant difference in germination. Under laboratory conditions, theseeds of L. apiculata ingested by birds presented astatistically greater percentage of germination than the seeds extracted manually.Under greenhouse conditions, seeds of M. planipes ingestedby birds did not present a statistically different germination percentage fromthose seeds extracted from the fruits. The seeds of M.planipes, and L. apiculata inside the intactfruits did not germinate. We conclude that birds do not affect the seedviability of any of the four species studied.