Native seed preferences of shrub-steppe rodents,birds and ants: the relationships of seed attributes and seed use

Summary This study established the preferences of shrubsteppe granivores among seeds of 6 common sagebrushsteppe plants and related the preferences observed to physical and nutritional attributes of the seeds. Seeds of big sagebrush (Artemisia tridentata), cheatgrass (Bromus tectorum), Indian ricegrass (Oryzopsis hymenoides), western wheatgrass (Pascopyrum smithii), bitterbrush (Purshia tridentata) and green needlegrass (Stipa viridula) were placed in groups of petri dishes designed such that seed removal could be ascribed to either diurnal vertebrates, nocturnal vertebrates or ants. Though absolute quantities of seeds removed varied among the 3 granivore classes, calculations of preference based on weights of each seed species removed by each granivore class indicated that all 3 ranked the seeds similarly. Preference hierarchies of the 3 granivore classes were highly positively correlated with both calories per seed and % soluble carbohydrate of the seeds. The first correlation supports a basic prediction of optimal foraging theory —that foragers should maximize energy intake per unit time spent foraging. Both correlations emphasize the role of seed nutritional qualities in granivore seed selectivity in that soluble carbohydrate is a water-efficient energy source and its percentage is a good indicator of the digestible energy available in a food item. A corollary experiment comparing granivore use of an exotic seed (millet [Panicum miliaceum]) and a preferred native seed (Oryzopsis) demonstrated a distinct preference for the exotic. Since millet seeds are particularly high in % soluble carbohydrate, this result reinforced the apparent value of this nutritional attribute as a predictor of granivore seed preference. Among many seed resource characteristics upon which granivore seed selectivity might operate, our results indicate that individual species' nutritional composition may be particularly important. Thus, inferences about seed selectivity and resource partitioning among arid-land granivores should be interpreted with caution, especially those based on experiments using seed introductions, since the influence of seed nutritional attributes has not been widely acknowledged heretofore.     

What do mice select for in seeds?

Summary Knowledge of the basis upon which granivores select seeds is crucial to the understanding of granivory. In this study the preferences of three rodent granivores among seeds of 11 plants from the semi-arid Karoo of South Africa were estimated, and related to the physical and chemical attributes of the seeds. Seed weights and calorific, moisture, protein, polyphenol, ash, lipid and silica contents were estimated and cell contents, soluble ash and soluble carbohydrate contents were derived from these values. These attributes were determined for both the intact seeds and the portion of the seed that is ingested by the mice. The efficiency with which mice ingested the seeds (in terms of time and mass) was recorded. All three mouse species ranked the seeds similarly, and the two species for which handling efficiency was measured did not differ in this regard. Preference hierachies were highly correlated with the rate of energy intake, as predicted by optimal foraging theory. There was no correlation between rodent preferences and the gross energy content of the seeds, emphasising the importance of measuring relevant parameters. The energy yield of the seeds calculated here, in conjunction with rodent population energy requirements and dietary data, may be used to estimate potential granivore impact on the seed production of the plant community.     

Spicing up restoration: can chili peppers improve restoration seeding by reducing seed predation?

Seed predation by rodents presents a significant barrier to native plant recruitment and can impede restoration seeding efforts. In nature, some plants contain secondary defense compounds that deter seed predators. If these natural defense compounds can be applied to unprotected seeds to inhibit rodent granivores, this approach could improve restoration seeding. Capsaicin is the active ingredient in chili pepper (Capsicum spp.) seeds that creates the burning sensation associated with human consumption of hot peppers. This compound has a similar effect on other mammals and is believed to have evolved as a deterrent to rodent seed predators. We used seed‐coating techniques to attach powder ground from Bhut Jolokia (Capsicum chinense) peppers to native plant seeds and evaluated the efficacy of these seed coatings for deterring rodent seed predation and enhancing native plant recruitment using laboratory and field experiments. Laboratory feeding trials demonstrated that native deer mice (Peromyscus maniculatus) consumed far fewer pepper‐coated seeds compared to untreated control seeds. Field seed‐addition experiments consistently demonstrated that rodent seed predation reduced native plant recruitment over the 4‐year study. Coating techniques used in the first 3 years were not persistent enough to reduce rodent seed predation effects on plant recruitment. However, a more persistent coating applied in conjunction with late‐winter sowing negated rodent seed predation effects on recruitment in year 4. Our results demonstrate that coating seeds with natural plant defense compounds may provide an effective, economical way to improve the efficacy of plant restoration by deterring seed predation by ubiquitous rodent granivores.     

Trophic and non-trophic pathways mediate apparent competition through post-dispersal seed predation in a Patagonian mixed forest

Post‐dispersal predation can be a major source of seed loss in temperate forests. Little is known, however, about how predator‐mediated indirect interactions such as apparent competition alter survival patterns of canopy tree seeds. Understorey plants may enhance tree seed predation by providing sheltered habitat to granivores (non‐trophic pathway). In addition, occurrence of different tree seeds in mixed patches may lead to short‐term apparent competition between seed types, because of the granivores’ foraging response to changes in food patch quality (trophic pathway). We hypothesised that understorey bamboo cover and mixing of seed species in food patches would both increase tree seed predation in a Nothofagus dombeyi?Austrocedrus chilensis forest in northern Patagonia, Argentina. Seed removal experiments were conducted for three consecutive years (2000–2002) differing in overall granivory rates. Seed patch encounter and seed removal rates were consistently higher for the larger and more nutritious Austrocedrus seeds than for the smaller Nothofagus seeds. Seed removal was greater beneath bamboo than in open areas. This apparent competition pathway was stronger in a low‐predation year (2000) than in high‐predation years (2001–2002), suggesting a shift in microhabitat use by rodents. Patch composition had a significant, though weaker, impact on seed survival across study years, whereas seed density per patch enhanced encounter rates but did not influence seed removal. Removal of the less‐preferred Nothofagus seeds increased in the presence of Austrocedrus seeds, but the reciprocal indirect effect was not observed. However, this non‐reciprocal apparent competition between seed species was only significant in the high‐predation years. Our study shows that granivore‐mediated indirect effects can arise through different interaction pathways, affecting seed survival patterns according to the predator's preference for alternative seed types. Moreover, results indicate that the occurrence and relative strength of trophic vs non‐trophic pathways of apparent competition may change under contrasting predation scenarios.     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

The strength of seeds and their destruction by granivorous insects

The influence of seed structure and strength on their destruction by granivores is central to understanding the dynamics of granivore-plant interactions. For up to nine seed species, the effects of seed size (cm³), mass (mg), density (mg/cm³) and coat strength (MPa) on the damage inflicted by three post-dispersal granivores (Harpalus pensylvanicus, Anisodactylus sanctaecrucis, and Gryllus pennsylvanicus) were evaluated. Seed destruction rates by G. pennsylvanicus were statistically unrelated to the size and toughness of the seeds. Seed densities significantly affected their destruction by A. sanctaecrucis and H. pensylvanicus, as did seed size, mass, and strength in H. pensylvanicus under choice conditions. The carabid beetles destroyed more of the small, denser seeds with stronger seed coats. The results show that different granivores are able to distinguish the structural strength and physical density of seeds as well as seed size. The relative ability of granivores to detect these seed characteristics offers a way in which diverse communities of post-dispersal insect granivores can persist within a single habitat. The authors redefine how the strength of biological structures should be evaluated in ecological studies, using guidelines commonplace in the field of engineering. Handling Editor: Stanislav Gorb The U.S. Government’s right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged     

Relationships among the seed rain,seed bank and vegetation of a Hawaiian forest

Abstract. Hawaiian ecosystems are prone to invasion by alien plant species. I compared the seed rain, seed bank, and vegetation of a native Hawaiian forest to examine the potential role that seed ecology plays in allowing alien species to invade native forest. Absolute cover of seed plants in the forest was 126 %, annual seed rain was 5 713 seeds m^-2 yr^-1, and the mean density of seedlings emerging from the seed bank averaged across four seasons was 1 020/m². The endemic tree Metrosideros polymorpha was the most abundant species in the vegetation, seed rain and winter seed bank. Overall, native seed plants comprised 95 % of the relative cover in the vegetation and 99 % of the seeds in the seed rain, but alien species comprised 67 % of the seeds in the seed bank. Alien species tended to form persistent seed banks while native species formed transient or pseudo-persistent seed banks. Dominance of the seed bank by alien species with persistent seed banks suggests that aliens are favorably placed to increase in abundance in the vegetation if the forest is disturbed.     

Fruit-seed relations in Hakea: serotinous species invest more dry matter in predispersal seed protection

Abstract The ability to maintain a canopy stored seedbank (serotiny) is characteristic of many woody genera inhabiting fire-prone environments. The relationship between level of serotiny for 94 Hakea (Proteaceae) species in southwestern Australia and follicle mass, density, three wall thicknesses and seed mass was investigated. Two species were non-serotinous (fruits open at maturity), 12 were weakly serotinous (majority of fruits open at maturity), 9 were moderately serotinous (fruits open within five years of maturity) and 71 were strongly serotinous (fruits still closed at least five years after reaching maturity). A positive relationship existed between the level of serotiny and follicle morphology. Strongly serotinous species were more likely to have heavier, woodier and thicker-walled follicles than non- and weakly serotinous species. Moderately/strongly serotinous species invested more energy (six times higher follicle:seed mass ratio) than non weakly serotinous species, consistent with increased protection of the seeds from granivores, pathogens, desiccation and/or heat. Recent work has shown that thicker fruit walls (strongly serotinous species) provide better insulation to seeds from heat, although the need to survive fire is just as critical for thin-walled, weakly serotinous species. Greater protection from granivores may provide a better explanation for the adaptive significance of dense, thick-walled serotinous follicles, as the opportunities for predispersal granivore damage are low among weakly serotinous species.     

Non-native mammals are weak candidates to substitute ecological function of native avian seed dispersers in an island ecosystem

Although prominent examples exist of non-native species causing substantial ecological harm, many have neutral or positive effects, including filling surrogate roles once performed by extinct native organisms. We tested the ecological roles of two non-native mammals as seed dispersers or seed predators in Guåhan, which, due to invasive brown tree snakes (Boiga irregularis), is devoid of native seed dispersers–birds and bats. We conducted feeding trials with captive rats (Rattus spp.), which are present but uncommon due to predation by snakes, and pigs (Sus scrofa), which are abundant. We examined if and how they interacted with common forest fruits. We then compared how any gut-passed or animal-handled seeds germinated compared to seeds left in whole fruit or depulped seeds. Rats and pigs interacted with most of the fruits and seeds (>80%) that they were fed. Of those, most seeds were destroyed—78% for rats and 90% for pigs, across both native and non-native plant species. Compared to seeds germinating within whole fruits, rats improved germination of the seeds that they handled without ingesting, while pigs diminished the germination of seeds that they handled. The small percentage of seeds (approximately 1.5% for rats and 5% for pigs) that survived gut passage germinated in higher proportions than those in whole fruits. Percentages of seed survival to germination are lower than found in similar studies with native avian frugivores. Our results indicate that pigs and rats have mixed effects on seeds, but are not suitable surrogates for native seed dispersers.     

Introduced galliforms as seed predators and dispersers in Hawaiian forests

In altered communities, novel species’ interactions may critically impact ecosystem functioning. One key ecosystem process, seed dispersal, often requires mutualistic interactions between frugivores and fruiting plants, and functional traits, such as seed width, may affect interaction outcomes. Forests of the Hawaiian Islands have experienced high species turnover, and introduced galliforms, the largest of the extant avian frugivores, consume fruit from both native and non-native plants. We investigated the roles of two galliform species as seed dispersers and seed predators in Hawaiian forests. Using captive Kalij Pheasants (Lophura leucomelanos) and Erckel’s Francolins (Pternistis erckelii), we measured the probability of seed survival during gut passage and seed germination following gut passage. We also examined which seeds are being dispersed in forests on the islands of O’ahu and Hawai’i. We found that galliforms are major seed predators for both native and non-native plants, with less than 5% of seeds surviving gut passage for all plants tested and in both bird species. Gut passage by Kalij Pheasants significantly reduced the probability of seeds germinating, especially for the native plants. Further, larger-seeded plants were both less likely to survive gut passage and to germinate. In the wild, galliforms dispersed native and non-native seeds at similar rates. Overall, our results suggest the introduced galliforms are a double-edged sword in conservation efforts; they may help reduce the spread of non-native plants, but they also destroy the seeds of some native plants. Broadly, we show mutualism breakdown may occur following high species turnover, and that functional traits can be useful for predicting outcomes from novel species’ interactions.

     

The transition from invasive species control to native species promotion and its dependence on seed density thresholds

Question: Does the seed density of invasive species affect establishment by native species in a bare ground context (following invasive species control efforts), and is it possible to promote transition to a native species dominated state by manipulating sowing density of the native community? Location: Experimental wetland basin in Chanhassen, Minnesota, USA. Methods: A mesocosm experiment investigated the influence of Phalaris arundinacea (invasive species) propagule pressure on establishment of native wet meadow species in the context of a newly restored wetland. Mesocosms were sown with P. arundinacea (0, 10, 50, 100, or 500 seeds/m²) and a mix of native species (3000 or 15000 seeds/m²). Results: When planted at densities > 100 seeds/m², P. arundinacea increased suppression of native species. Also, high native seed density suppressed P. arundinacea biomass production. This effect was more pronounced when P. arundinacea seed density was high (> 100 seeds/m²), but high native seed density (15000 seeds/m²) did not suppress recruitment of P. arundinacea from seed. Conclusions: The transition from post‐control bare ground (a common result of efforts to control invasive species) to native species establishment depends on both native species and invader seed density. These results suggest that a threshold of P. arundinacea propagule pressure exists, beyond which transition to a native community is less likely without management intervention. P. arundinacea can establish in the presence of a newly developing native plant community, even at very low densities of P. arundinacea seed. Invader control (following initial site clearing efforts) is essential to native species establishment.     

Granivory in California sage scrub: implications for common plant invaders and ecosystem conservation

Seed predation may influence community assembly and invasion dynamics when seed predators preferentially select some seed species over others. However, the role of different seed predators and their preferences for seeds in multiple ecological contexts, including the endangered California sage scrub ecosystem, remain unresolved making predictions about which processes limit or promote invasions difficult. In addition, selection criteria, specifically how seed size and biogeographic origin (native versus invasive) influences selection, requires further research across multiple granivore guilds. We examined seed predator preference for common native and invasive seeds across small and large seed classes using seed dish experiments with motion-sensor video observation. We also quantified the relative importance of ants, birds, and mammals as seed predators in sage scrub. Community-wide, granivores preferred the small-seeded invasive mustard Brassica nigra, avoided the large-seeded invasive grass Bromus rubens and the native large-seeded shrub Encelia californica, and did not show significant selection or avoidance for the small-seeded native shrub Salvia apiana. Birds, notably California towhees (Melozone crissalis), were the most frequent granivore while rodents and ants rarely removed seeds. This study reveals that birds are important seed predators in California sage scrub and have the potential to contribute to biotic resistance to mustard invasions (Brassica nigra and potentially others).

     

Seed storage: maintaining seed viability and vigor for restoration use

Effective seed storage after sourcing (harvesting or purchasing) is critical to restoration practitioners and native seed producers, as it is key to maintaining seed viability. Inadequate seed storage can lead to a waste of both natural and economic resources when seeds of poor quality are sown. When working with native species with unknown storage behavior, general assumptions can be made based on studies on related species, and standard practices may be applied with caution; however, an investigation should be conducted to understand if specific storage requirements are needed and for how long seeds can be stored before they lose significant viability. In this paper of the Special Issue Standards for Native Seeds in Ecological Restoration, we provide an overview of the key concepts in seed storage and the steps to take for effective storage of native seeds for restoration use.     

A multi-species assessment of post-dispersal seed predation in the central Chilean Andes

BACKGROUND AND AIMS: Post-dispersal seed predation in alpine communities has received little attention despite evidence that seeds removed by granivores can decrease plant recruitment into ecosystems. Moreover, few studies have assessed the effects of removal of seeds of a range of species after dispersal on the seeds remaining in ecosystems. A comparison was made of the magnitude of seed removal by ants and birds of nine different shrubby-, herbaceous- and cushion-plant species in the central Chilean Andes in order to assess the interactions between birds, ants and wind, and the types of seeds. METHODS: A total of 324 soil-covered plates, each containing 50 seeds of one species, were placed in the field at an altitude of 2700 m and assigned to one of four treatments: control, exclusion of ants, birds, and both. The design also allowed the effects of wind to be assessed. Seed removal from plates was monitored over 20 d. KEY RESULTS: Mean accumulative seed removal by granivores averaged over all nine species combined was 25%. However, large differences between species were evident, with limited seed removal (3-11%) in three herbaceous species (Alstroemeria pallida, Sisyrinchium arenarium, Pozoa coriacea), moderate (18-33%) in five species, including a shrub (Chuquiraga oppositifolia), two herbs (Taraxacum officinale, Rhodophiala rhodolirion), and two cushion-plants (Laretia acaulis, Azorella monantha), and substantial (78%) in the shrub Anarthrophyllum cumingii. The magnitudes of losses caused by birds compared with ants did not differ for the majority of species, although removal by birds was greater than by ants in A. cumingii, and smaller for C. oppositifolia. CONCLUSIONS: Post-dispersal seed removal is shown to be an important cause of decreased potential plant species recruitment into alpine ecosystems. The substantial differences in the magnitude of seed losses to ants and birds demonstrate the need for evaluation of seed removal on a wide range of species in any given ecosystem.     

The role of avian ‘seed predators’ as seed dispersers

Seed dispersal is a central process in plant ecology with consequences for species composition and habitat structure. Some bird species are known to disperse the seeds they ingest, whereas others, termed ‘seed predators’, digest them and apparently play no part in dispersal, but it is not clear if these are discrete strategies or simply the ends of a continuum. We assessed dispersal effectiveness by combining analysis of faecal samples and bird density. The droppings of seed dispersers contained more entire seeds than those of typical seed predators, but over a quarter of the droppings of seed predators contained whole seeds. This effect was further magnified when bird density was taken into account, and was driven largely by one frequent interaction: the Chaffinch Fringilla coelebs, a typical seed predator and the most abundant bird species in the area and dispersed seeds of Leycesteria formosa, a non‐native plant with berry‐like fruits. These results suggest the existence of a continuum between seed predators and seed dispersers.     

Differential weed seed removal in dryland cereals

Weeds persist in rain-fed cereal fields in NE Spain, despite intense herbicide use and high seed removal rates by granivorous harvester ants. Herbicide resistance is involved, but certain weed species also appear to escape seed removal by granivores. To identify the mechanisms involved, we measured seed removal rates (three fields in 2010) and the timing of seed shed (one field in 2009 and three fields in 2010) and used an existing model, which integrates short-term rates of seed shed, burial and removal, to estimate long-term seed removal rates. Averaged over years, fields and weed species, the long-term seed removal rate was estimated at 72?% (range 46?C100?%). Fifteen to 25?% of the seeds of Bromus diandrus avoided removal by being less attractive (low removal rates), and another 0?C29?% escaped through crop harvest, which made seeds inaccessible to granivores. Similarly, 20?C32?% of the Papaver rhoeas seeds escaped through crop harvest, while another 13?C17?% escaped by burial into the soil (small seed size). Other species, such as Galium spurium or Diplotaxis erucoides, had no means of avoiding seed removal by harvester ants. In particular, the more troublesome weeds, such as B. diandrus, P. rhoeas and L.?rigidum, combined herbicide resistance or tolerance with avoidance mechanisms against granivory.     

Functional extinction of a desert rodent: implications for seed fate and vegetation dynamics

Population declines of once‐abundant species have often preceded understanding of their roles within ecosystems. Consequently, important drivers of environmental change may remain undiagnosed because we simply do not know how species that are now rare or extinct shaped ecosystems in the past. Australia's desert rodents are thought to have little numerical impact on seed fate and vegetation recruitment when compared with ants or with desert rodents on other continents. However most research on granivory by Australian desert rodents has occurred in areas where rodents were rare or functionally extinct. Here we ask if the paradigm that rodents are relatively un‐important granivores in Australian deserts is an artefact of their historical decline. In the Strzelecki Desert, the endangered rodent, Notomys fuscus is rare where introduced mesopredators are abundant but common where dingoes (an apex predator) suppress mesopredator populations. We used foraging trays to compare rates of seed removal for a common shrub (Dodonaea viscosa angustissima hopbush) between areas where N. fuscus, hopbush shrubs and their seedlings were rare and common and found that seed removal was consistently higher where rodents were common and hopbush rare. By excluding ants and rodents from foraging trays we show that ants removed more seeds than rodents where rodents were rare but rodents removed far more seeds than ants where rodents were common. By manipulating rodents’ access to the soil seed‐bank we show that hopbush seeds persisted in greater numbers where rodents were excluded than where they had access. Our results support the hypothesis that granivory by rodents may once have been a far more important process influencing the fate of seeds and shaping plant communities in arid Australia and suggest that dingo extirpation has cascading effects on shrub seeds. Our study highlights that functional extinction of rodents may be an under‐appreciated driver of vegetation change.     

Foraging,food selection and worker size in the seed harvesting ant <Emphasis Type="Italic">Messor bouvieri</Emphasis>

Interactions among granivores and seeds depend on the foraging behaviour and morphology of the granivores and on the attributes and availability of seeds. We investigated seed selection by the seed harvesting ant Messor bouvieri in three adjacent plant communities in Spain by relating the harvested seeds to those in the seed rain. Preference was positively correlated with seed size and abundance which accounted for 43% and 20% of the variance respectively. Contrary to predictions of central place foraging theory, the size of seeds harvested did not increase with distance from the nest. Inclusion of a less-preferred item in the diet was more strongly related to the abundance of more-preferred items (60% of the variance) than the abundance of the less-preferred item (14% of the variance). Worker size accounted for 20–30% of the variance in the size of harvested seeds, although small workers did not appear to be constrained by load size for the range of seeds available. The body size of ants was significantly larger in the community with the greatest proportion of large seeds, although this was not due to their ability to carry larger loads or due to the greater force required to crush these seeds. The strong preference of M. bouvieri for large seeds may have important consequences for the plant communities in which they forage. Received: 30 November 1999 / Accepted: 14 April 2000     

Mechanisms affecting the fate of Prosopis flexuosa (Fabaceae,Mimosoideae) seeds during early secondary dispersal in the Monte Desert,Argentina

Abstract The fate of seeds during secondary dispersal is largely unknown for most species in most ecosystems. This paper deals with sources of seed output of Prosopis flexuosa D.C. (Fabaceae, Mimosoideae) from the surface soil seed‐bank. Prosopis flexuosa is the main tree species in the central Monte Desert, Argentina. In spite of occasional high fruit production, P. flexuosa seeds are not usually found in the soil, suggesting that this species does not form a persistent soil seed‐bank. The magnitude of removal by animals and germination of P. flexuosa seeds was experimentally analysed during the first stage of secondary dispersal (early autumn). The proportion of seeds removed by granivores was assessed by offering different types of diaspores: free seeds, seeds inside intact endocarps, pod segments consisting of 2–3 seeds, and seeds from faeces of one herbivorous hystricognath rodent, the mara (Dolichotis patagonum). The proportion of seeds lost through germination was measured for seeds inside intact endocarps, seeds inside artificially broken endocarps, and free seeds. Removal by ants and mammals is the main factor limiting the formation of a persistent soil seed‐bank of P. flexuosa: >90% of the offered seeds were removed within 24 h of exposure to granivores in three of four treatments. Seeds from the faeces of maras, on the other hand, were less vulnerable to granivory than were other types of diaspores. These results suggest that herbivory might be an indirect mechanism promoting seed longevity in the soil (and likely germination) by discouraging granivore attack. On the other hand, germination did not seem to have an important postdispersal impact on the persistence of P. flexuosa seeds in the soil. Both direct and indirect interactions between vertebrate herbivores and plants may foster P. flexuosa's seed germination in some South American arid zones.     

Small mammals as potential seed dispersers in New Zealand

Weed invasion success is strongly influenced by availability of seed dispersal vectors, which may include animals. We examined the potential of several small introduced mammals (mice, kiore, ship rats and possums) to disperse germinable seeds in New Zealand. Captive animals were fed fleshy fruit of weeds (Berberis glaucocarpa, Cotoneaster spp., Crataegus monogyna, Ilex aquifolium, Leycesteria formosa, Ligustrum sinense, Lonicera japonica, Passiflora mollissima, Pyracantha angustifolia, Sorbus hupehensis) and native species (Coprosma spp., Prumnopitys ferruginea and Solanum aviculare). We recorded the percentage of fruit consumed, seed ingested and gut passage time. Faeces were collected and the seeds extracted and tested for germination potential in an unheated glasshouse (two weed species) or under controlled conditions (11 species). The smallest rodents (mice and kiore) generally destroyed all seeds eaten. Large numbers of viable seeds of the small‐seeded (<1 mg) species, L. formosa and S. aviculare, passed through ship rats. Possums consumed the seeds of all adventive and native fruits except P. ferruginea. The proportion of seeds recovered intact from possum faeces varied with plant species and ranged from 6 to 83%. The time required for 50% of all seeds to be passed by possums ranged from 2.5 to 5.5 days with an average of 3.7 days, and was generally unrelated to simple fruit parameters such as percentage pulp and moisture content. For seeds where germination also occurred in the uneaten controls, the germination of seed from possums ranged from 3 to 78%. Germination was mostly lower in seeds from possums than in the controls, where differences were significant. Possums have major potential to disperse a wide range of fleshy fruit‐producing native and introduced plant species. Ship rats have the potential to disperse those with very small seeds.