Spatial Variation in the Strength of a Trophic Cascade Involving Ruellia nudiflora (Acanthaceae), an Insect Seed Predator and Associated Parasitoid Fauna in Mexico

Spatial variation in the strength of herbivore top-down control represents an important source of variation in plant fitness measures and community structure and function. By measuring seed predator (larvae of a Noctuid moth) and parasitoid impacts on Ruellia nudiflora across a broad spatial scale in Yucatan (Mexico), this study addressed the following: (1) to what extent does seed predator and parasitoid attack intensity associated with R. nudiflora vary spatially? (2) Does parasitoid attack result in a positive indirect effect on the plant, and does the intensity of this effect vary spatially? During the peak of fruit production (late June–early July) of 2005, we collected fruits from 21 R. nudiflora populations and grouped them into four regions: center, east, north and south. For each fruit we recorded: observed seed number, number of seeds eaten, seed predator presence, parasitoid presence and number of seeds ‘saved’ by parasitoids. Seed predators attacked ca 30 percent of fruits/plant on average, while parasitoids were found in 24 percent of seed predator-attacked fruits. Results indicated spatial variation in seed predator and parasitoid attack levels; interestingly, a contrasting spatial gradient of attack intensity was observed: populations/regions with greatest parasitoid attack levels usually had the lowest seed predator attack levels and vice versa, suggesting top-down control of parasitoids on seed predators. We observed a weak overall indirect impact of parasitoids on R. nudiflora (4% seeds ‘saved’ on average), which nonetheless varied strongly across populations (e.g., close to 14% seeds saved at one population). Findings indicate a geographical structuring of interaction strengths across populations, as well as spatial variation in the strength of parasitoid cascading effects on plant reproduction.     

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.     

Responses of pre‐dispersal seed predators to sequential flowering of Dipterocarps in Malaysia

Many species of Dipterocarpaceae and other plant families reproduce synchronously at irregular, multi‐year intervals in Southeast Asian forests. These community‐wide general flowering events are thought to facilitate seed survival through satiation of generalist seed predators. During a general flowering event, closely related Shorea species (Dipterocarpaceae) stagger their flowering times by several weeks, which may minimize cross pollination and interspecific competition for pollinators. Generalist, pre‐dispersal seed predators might also track flowering hosts and influence predator satiation. We addressed the question of whether pre‐dispersal seed predation differed between early and late flowering Shorea species by monitoring flowering, fruiting and seed predation intensity over two general flowering events at the Pasoh Research Forest, Malaysia. Pre‐dispersal insect seed predators killed up to 63 percent of developing seeds, with Nanophyes shoreae, a weevil that feeds on immature seeds being the most important predator for all Shorea species. This weevil caused significantly greater pre‐dispersal seed predation in earlier flowering species. Long larval development time precluded oviposition by adults that emerged from the earliest flowering Shorea on the final flowering Shorea. In contrast, larvae of weevils that feed on mature seeds before seed dispersal (Alcidodes spp.), appeared in seeds of all Shorea species almost simultaneously. We conclude that general flowering events have the potential to satiate post‐dispersal seed predators and pre‐dispersal seed predators of mature fruit, but are less effective at satiating pre‐dispersal predators of immature fruit attacking early flowering species.     

Consequences of the timing of seed release of Erythronium americanum (Liliaceae), a deciduous forest myrmecochore

Myrmecochores are plants with seeds adapted for ant dispersal. This specialized dispersal syndrome may provide Erythronium americanum seeds with protection from predators within the eastern deciduous forests. To determine the adaptive significance of myrmecochory in E. americanum, seed removal rates and seed predation in relation to seed release date and location along the Potomac River in Langley, Virginia, were examined. The number of seeds removed from four exclosure treatments were monitored two times in 1992 and three times in 1993 within floodplain and hilltop populations of E. americanum. Overall, seed removal was greatest from control depots, and E. americanum seeds were removed at nearly the same rate from predator-exclusion depots, indicating that removal from open depots is largely due to ant removal. Ants removed significantly more seeds than predators in the first 48 h of seed exposure and could potentially remove all E. americanum seeds before nightfall. Aphaenogaster rudis was identified as the primary disperser of E. americanum. Seeds placed in depots after the natural seed release period were discovered more quickly and removed by ants at a significantly higher rate than seeds released at the natural date. These results suggest that ant dispersal of E. americanum seeds reduces the likelihood of seed predation.     

Invertebrate Seed Predators are not all the Same: Seed Predation by Bruchine and Scolytine Beetles Affects Palm Recruitment in Different Ways

Because most invertebrate seed predators are host‐specific, they are usually expected to produce Janzen–Connell patterns. This expectation was fulfilled for Astrocaryum but not for Allagoptera, depending on the effects of bruchine and scolytine predators on the seeds of these palms. Thus, the mere existence of invertebrate predation is not sufficient for generating Janzen–Connell; what matters is seed mortality, which varies between predators and between plant species for the same predator.     

Intensity of pre‐dispersal seed predation in the invasive legume Leucaena leucocephala is limited by the duration of pod retention

Abstract The intensity of seed predation the invasive tropical legume Leucaena leucocephala by the bruchid Acanthoscelides macropthalmus was investigated in south‐eastern Queensland, Australia. The number of seeds damaged by A. macropthalmus as a proportion of total seeds available was found to increase the longer the pods remained on the tree. Seed predation ranged from a mean of 10.75% of seeds on pods that remained on the plant for 1 month and increased to 53.54% for pods that remained of the plant for 4 months. The low bruchid populations at high pod densities results in ‘predator satiation’. However, pods dehisce over time and the proportion of pods available over time to the bruchid correspondingly declines. By the time bruchid densities build up, most pods have dehisced and the seeds consequently escape predation. As a result the number of seeds lost to bruchid damage increases only marginally over time. Despite the levels of seed predation observed over the course of the study, the number of seeds in the soil seedbank almost doubled over time increasing from 8.5 seeds m^?3 to 15.5 seeds m^?3 over a 4‐month period. Levels of seed predation and addition of seeds to the soil seedbank were not correlated. The taxonomic (subspecies) status and apparency of host plants as measured by plant and patch traits (average plant height, density of podding plants and patch size) did not influence levels of seed predation. Pre‐dispersal seed predation studies need to take into account the pod/seed retention behaviour of the plant. The ability of the bruchid to regulate the invasiveness of Leucaena through influencing its demography is likely to be diminished if the insect populations cannot increase rapidly enough to use the seeds before pod dehiscence.     

Habitat effects on second-order predation of the seed predator Harpalus rufipes and implications for weed seedbank management

Seed predators provide a valuable ecosystem service to farmers by reducing densities of weed seeds, and, in turn, densities of weed seedlings they must manage. The predominant invertebrate weed seed predator in Maine, USA, agroecosystems is the carabid beetle Harpalus rufipes DeGeer. Pitfall trapping has shown that H. rufipes prefers sites with vegetative cover to fallow sites, preference speculated to be driven by predator avoidance behavior. To test this hypothesis, ‘second-order predation assays’ were developed, in which live H. rufipes prey were presented to second-order predators. Field experiments were conducted to determine foremost if H. rufipes was subject to second-order predation, and secondly, whether (a) vegetative cover affords H. rufipes protection from second-order predators, and (b) high rates of second-order predation correspond with decreased invertebrate seed predation rates. Two 72-h experiments were conducted (mid August and September 2012) at crop and non-crop sites across a 28 ha diversified farm in Stillwater, ME, USA.Second-order predation was 2.8% per day. Based on images from motion-sensing cameras, H. rufipes’ predators included birds and small mammals. Neither a relationship between second-order predation and vegetative treatment, nor an empirical relationship between second-order predation and invertebrate seed predation were detected. However, a simulation model predicted that 2.8% per day second-order predation could increase the number of seeds entering the seedbank by more than 17% annually. Additionally, complex habitats supported higher rates of second-order predation than did simple habitats.     

Comparing methods of weed seed exposure to predators

Several methods of seed exposure are used in seed predation studies, but how these methods influence the results remains poorly studied. In this article, two commonly used methods of seed exposure – seed cards and plasticine trays – were compared in the field and in the laboratory using three species of weed seeds. In the field, cards or trays with seeds were exposed either with or without cages to keep vertebrates out and either with or without impermeable roofs to provide protection from the weather. The overall seed retrieval from the control stands varied significantly between the methods of seed exposure, roof treatment and the species of seeds, and the scatter in the seed retrieval increased or decreased monotonically with the temperature or precipitation for some species of seeds. This indicates that the controls were more or less relevant depending on the weather conditions and species. The seed removal from cards varied between the species of seeds. The effect of exclosure cages indicated that invertebrates were the dominant seed predators of Capsella bursa‐pastoris, while in Poa annua and Stellaria media both vertebrates and invertebrates were important. Higher seed removal from plasticine trays compared to seed cards was found for all three species of seeds, and placing roofs over the seeds affected seed removal in C. bursa‐pastoris and P. annua. In the laboratory, seed consumption varied with the method of exposure in 10 out of 12 combinations of seed species and predator (two species of carabids and two species of isopods). The overall tendency was the opposite of the field observations: seed consumption was higher for seeds on filter paper and cards compared to seeds on tin trays. We conclude that seed cards are more convenient for short term studies in the field, while tin trays are useful in laboratory multi‐choice experiments. To measure the realistic consumption of invertebrate predators in the laboratory, filter paper seems to be the best option as it does not present an obstacle to predators eating the seed. Using roofs introduced further bias regarding the estimates of seed removal in the field and thus should be avoided.     

The biology of seed discrimination and its role in shaping the foraging ecology of carabids: A review

Species of carabid (ground) beetles are among the most important postdispersal weed seed predators in temperate arable lands. Field studies have shown that carabid beetles can remove upwards of 65%–90% of specific weed seeds shed in arable fields each year. Such data do not explain how and why carabid predators go after weed seeds, however. It remains to be proven that weed seed predation by carabids is a genuine ecological interaction driven by certain ecological factors or functional traits that determine interaction strength and power predation dynamics, bringing about therefore a natural regulation of weed populations. Along these lines, this review ties together the lines of evidence around weed seed predation by carabid predators. Chemoperception rather than vision seems to be the primary sensory mechanism guiding seed detection and seed selection decisions in carabid weed seed predators. Selection of weed seeds by carabid seed predators appears directed rather than random. Yet, the nature of the chemical cues mediating detection of different seed species and identification of the suitable seed type among them remains unknown. Selection of certain types of weed seeds cannot be predicted based on seed chemistry per se in all cases, however. Rather, seed selection decisions are ruled by sophisticated behavioral mechanisms comprising the assessment of both chemical and physical characteristics of the seed. The ultimate selection of certain weed seed types is determined by how the chemical and physical properties of the seed match with the functional traits of the predator in terms of seed handling ability. Seed density, in addition to chemical and physical seed traits, is also an important factor that is likely to shape seed selection decisions in carabid weed seed predators. Carabid responses to seed density are rather complex as they are influenced not only by seed numbers but also by trait‐based suitability ranks of the different seed types available in the environment.     

Testing the quick meal hypothesis: The effect of pulp on hoarding and seed predation of Hymenaea courbaril by red‐rumped agoutis (Dasyprocta leporina)

Abstract: Red‐rumped agoutis (Dasyprocta leporina) are important seed dispersers/predators of Neotropical large‐seeded plants. Several species of seeds cached by agoutis have an edible reward, in contrast to temperate rodent‐dispersed diaspores. The quick meal hypothesis states that the presence of a reward such as edible pulp will enhance the efficiency of rodents as seed disperses by satiating the animal and, consequently, reducing seed predation and enhancing hoarding. In this study, this hypothesis was tested using as the reference system the pulp and seeds of Hymenaea courbaril. Seeds with and without pulp were offered to agoutis and the behaviour of each individual was recorded. Since the probability of predation and hoarding were complementary, we used the probability of predation. The proportion of agoutis that preyed on at least one seed was similar for seeds with (42.8% of individuals) and without (40.0% of individuals) pulp. In agoutis that preyed upon at least one seed, the probability that they killed a seed did not differ between seeds with (0.17 ± 0.03) and without (0.20 ± 0.08) pulp. Hence, these results do not support the ‘quick meal hypothesis’.     

Seed predation of Copaifera langsdorffii (Fabaceae): a tropical tree with supra‐annual fruiting

Seed predation is an important ecological and evolutionary force that directly affects the distribution of plant species. Copaifera langsdorffii is a tropical tree species with supra‐annual fruiting, which has its seeds predated by a specialist endogenous insect (Rynochenus brevicollis: Curculionidae) in the Brazilian savanna. Three hypotheses were addressed: (i) the predator satiation hypothesis, (ii) the resource concentration hypothesis and (iii) the larger seed predation hypothesis. A total of 112 individual C. langsdorffii were monitored monthly from January to August during four consecutive years (from 2008 to 2011) to determine the presence of fruits on each plant. All trees produced fruits in the year 2008, whereas none of them produced flowers or fruits in 2009 or 2010. Moreover, only 65 individuals (58%) marked in 2008 produced fruits in 2011. The number of fruits per plant was approximately 21% greater in 2008 than in 2011, while the percentage of seed predation was 76% greater in 2011, thereby supporting the predator satiation hypothesis. The percentage of seeds predated was not affected by the number of fruits per plant. Therefore, our data did not support the resource concentration hypothesis. Plants producing large seeds experienced more seed predation by R. brevicollis, supporting the larger seed predation hypothesis. In addition, we also observed a positive relationship between seed volume and adult R. brevicollis weight. This study demonstrates the importance of supra‐annual fruiting for increasing survivorship of C. langsdorffii seeds both at the individual and the population level, and suggests that seed predators select plants producing large seeds as a way of increasing the number of offspring.     

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.     

Seed predator deterrence by seed-carrying ants in a dyszoochorous plant, <Emphasis Type="Italic">Chamaesyce maculata</Emphasis> L. Small (Euphorbiaceae)

Seeds are often carried by omnivorous ants even if they do not carry elaiosomes. Although many seeds carried by ants are consumed, both seeds abandoned during the seed carrying and leftover seeds are consequently dispersed (dyszoochory). These non-myrmecochorous seeds do not necessarily attract ants quickly. Therefore, these seeds often seem to be exposed to the danger of consumption by pre-dispersal seed predators. We propose the hypotheses, “seed predator deterrence hypothesis” that plants may benefit from seed-carrying ants if they deter seed predators from visiting plants, and seed-carrying ants may play additional roles in plant reproductive success, besides dyszoochory by ants. To test the hypotheses, we investigated the abundance of seed-carrying ants of the species Tetramorium tsushimae Linnaeus and Pheidole noda Smith F., and of the seed predatory stinkbug, Nysius plebeius Distat, on the spotted sandmat, Chamaesyce maculata L. Small, of which the seeds have no elaiosomes but are consumed by both ants and bugs. In the field, ants and stinkbugs seldom encountered each other on the plant. The number of stinkbugs beneath the plants with ants was smaller than that beneath the plants without ants. In laboratory experiments, the number of stinkbugs on the shoot was smaller when ants were present than when they were absent. These results might support the seed predator deterrence hypothesis: the probability of seed predation by stinkbugs seems to be reduced by the ant visits on plants and/or the existence of ants beneath the plants. This study highlights a new ant–plant interaction in seed dispersal by ants.     

Mammal Abundances and Seed Traits Control the Seed Dispersal and Predation Roles of Terrestrial Mammals in a Costa Rican Forest

In Neotropical forests, mammals act as seed dispersers and predators. To prevent seed predation and promote dispersal, seeds exhibit physical or chemical defenses. Collared peccaries (Pecari tajacu) cannot eat some hard seeds, but can digest chemically defended seeds. Central American agoutis (Dasyprocta punctata) gnaw through hard‐walled seeds, but cannot consume chemically defended seeds. The objectives of this study were to determine relative peccary and agouti abundances within a lowland forest in Costa Rica and to assess how these two mammals affect the survival of large seeds that have no defenses (Iriartea deltoidea, Socratea exorrhiza), physical defenses (Astrocaryum alatum, Dipteryx panamensis), or chemical defenses (Mucuna holtonii) against seed predators. Mammal abundances were determined over 3 yrs from open‐access motion‐detecting camera trap photos. Using semi‐permeable mammal exclosures and thread‐marked seeds, predation and dispersal by mammals for each seed species were quantified. Abundances of peccaries were up to six times higher than those of agoutis over 3 yrs, but neither peccary nor agouti abundances differed across years. Seeds of A. alatum were predominantly dispersed by peccaries, which did not eat A. alatum seeds, whereas non‐defended and chemically defended seeds suffered high levels of predation, mostly by peccaries. Agoutis did not eat M. holtonii seeds. Peccaries and agoutis did not differ in the distances they dispersed seeds. This study shows that seed fates are contingent upon many factors such as seed defenses, frugivore–granivore abundances, and seed‐handling capabilities. Mammal–seed interactions are complex; the outcomes of these interactions depend on the inherent characteristics of seeds and their potential dispersers.     

Predispersal seed predation in Bartsia alpina

Summary A northern Swedish population of Bartsia alpina, an arctic-alpine perennial herb, was found to suffer high levels of predispersal seed predation by larvae of two insect species, both specialists on rhinanthoid Scrophulariaceae hosts. The primary predator is Aethes deutschiana (Lepidoptera-Tortricidae), the host of which was previously unknown. The other predator is Gimnomera dorsata (Diptera-Scatophagidae), which is basically a Pedicularis specialist. Both predators are attacked by larvae of Scambus brevicorais (Hymenoptera-Parasitica-Ichneumonidae). Total predation pressure was more or less constant during 1985–1987, but in 1988 the level was doubled, the possible reasons of which are discussed. Large inflorescences of B. alpina suffer significantly higher predation pressures than small ones. It is shown that predation is most intense in the middle of the inflorescences. The same floral nodes are known to produce more selfed seeds than distal and basal nodes. Seed predation in B. alpina thus results in an increased proportion of outcrossed seeds entering the seed pool. Selection pressures on host plant and predator fauna are discussed.     

Do Meyer’s Parrots Poicephalus meyeri benefit pollination and seed dispersal of plants in the Okavango Delta,Botswana?

Recent studies of new world parrots repeatedly document, with few exceptions, that parrots are wasteful and destructive predispersal seed predators that are unlikely to contribute towards pollination and seed dispersal. Few detailed studies, however, have assessed the contribution of African parrots to forest ecology by quantifying the potential net benefit of seed and flower predation by parrots for most tree species in their diet. Due to the incidence of pollen on the heads of Meyer’s Parrots when feeding on Leguminosae flowers and the dispersal of viable seeds to the ground during seed predation, we compared destruction rates, when feeding on pods, fruits and flowers, with dispersal rates of viable seeds to the ground and frequency of head contact with reproductive apparatus to estimate net benefit from Meyer’s Parrot feeding activity. Meyer’s Parrots were not implicated in endo‐ or epizoochory, but they dropped uneaten fruit pulp and seeds to the ground during feeding bouts, thus providing ripe, undamaged seeds to secondary seed dispersers. This link with forest recruitment was weak, as all tree species utilized by Meyer’s Parrots either had more significant primary dispersal agents or were primarily wind‐dispersed. In most cases, the negative effect of seed predation outweighed any positive effects in terms of dispersal, whereby almost three times more seeds were consumed or destroyed than were dispersed to the ground. Significantly, only Sclerocarya birrea caffra recorded marginal net dispersal benefit from utilization by Meyer’s Parrots. Due to low relative resource abundance and high destruction rate, feeding activity on Berchemia discolor may be significant enough to influence its spatial distribution and abundance. Utilization of flowers of Kigelia africana and Adansonia digitata by parrots likely had a significant negative impact on pollination. Feeding on Acacia nigrescens flowers, however, was potentially advantageous to their pollination. We conclude that Poicephalus parrots are net consumers of ripe, undamaged seeds and flowers, thus having an overall negative impact on forest recruitment in subtropical Africa.     

Are rodents a source of biotic resistance to tree invasion in Pampean grasslands? Tree seed consumption under different conditions

Biotic resistance has been invoked as a major barrier to woody species invasion, although the role of resident generalist consumers and their interaction with seed availability in a local community has received little attention. We assessed tree seed consumption by rodents under two different scenarios: (i) We documented in field spatio‐temporal patterns of seed predation by native rodents on two exotic tree species, Gleditsia triacanthos or ‘honey locust’ and Robinia pseudoacacia or ‘white locust’ (family Leguminosae), in five grassland habitats of the Inland Pampa, Argentina. (ii) We conducted laboratory feeding trials to evaluate tree seed consumption in the presence (cafeteria‐style feeding trials) and in the absence (non‐choice feeding trials) of alternative food supplies. Seed predation was generally higher for Robinia than for Gleditsia seeds, both in field and laboratory conditions. For both tree species, seed predation varied between habitats and seasons and was higher in the native tussock grassland than in the remaining studied communities, whereas the crop field showed the lowest levels of consumption along with the absence of captured rodents. Seed consumption of Gleditsia and Robinia among the four grassland communities (which did not differ in rodent abundance) was negatively associated with the availability of alternative food. Laboratory feeding trials showed a higher consumption of Gleditsia seeds in the non‐choice than in the cafeteria‐style feeding trials, while the consumption of Robinia seeds did not differ in the absence or presence of alternative seeds. These patterns indicate that the contribution of resident granivores to invasion resistance might depend on colonizer species identity, recipient community type and season of the year. We suggest that rodent preferences for different invader seeds will interact with the availability of alternative food in the local habitat in influencing the amount of predator‐mediated biotic resistance to invasion.     

Unexpected relationships and valuable mistakes: non‐myrmecochorous Prosopis dispersed by messy leafcutting ants in harvesting their seeds

Abstract Ants generally disperse seeds while feeding on fruits or structures attached to the seed. Seed dispersal as a by‐product of seed predation (dyszoochory) was recognized in specialized harvester ants, but not in ants predating seeds opportunistically. Leafcutting ants are the main herbivores in much of the Neotropics, and they have been reported to remove fruits and seeds, but their role as seed predators and dispersers has not been acknowledged. Prosopis flexuosa D.C. (Fabaceae, Mimosoideae) is the most abundant tree species in the central Monte Desert, Argentina, and it is likely to depend on secondary animal dispersal. Mammalian frugivores are usually considered its main dispersers, but the opportunity for dispersal may be small since the removal of fruits and seeds by seed predators is very intense. The objective of this study was to identify which ant species interact with P. flexuosa fruits and to evaluate their relative importance as seed predators and dispersers. In a field experiment, whole and segmented pods were offered and several ant species exploiting the fruits were identified. Additionally, all pod segments remaining around nests of the three ant species able to remove them (the leafcutters Acromyrmex lobicornis Emery and Acromyrmex striatus Roger, and Pheidole bergi Mayr) were examined during and after the P. flexuosa primary dispersal season. Up to 753 pod segments and 90 sound seeds were found accumulated in a circle of 1 m radius over nests of A. lobicornis, and even more in an examined trail. Acromyrmex striatus left a smaller proportion of sound seeds and P. bergi left a smaller number of pod segments. All tendencies were similar during shorter known periods of accumulation. Leafcutting ants are acting as important seed predators, and ‘by mistake’ may be dispersing a key non‐myrmecochorous tree. This is an unexplored path in the seed dispersal cycle of P. flexuosa that challenges the tendency to predict interactions based on classifications made with other goals.     

Adaptive advantages of myrmecochory: the predator-avoidance hypothesis tested over a wide geographic range

The predator‐avoidance hypothesis states that once released from the parent plant, myrmecochorous seeds are rapidly taken by ants to their nests, where they are protected from predators. Previous studies conducted to test this hypothesis have frequently neglected two major aspects necessary for its verification: 1) the influence of processes acting after the seed release and 2) the spatial evenness of such processes. Thus, large‐scale variations in the mechanisms acting beyond seed release, and possibly influencing seed escape from predators, remain poorly documented. Here, we present the results of a post‐dispersal seed‐removal experiment on the myrmecochorous herb Helleborus foetidus, aimed at verifing the predator‐avoidance hypothesis by considering two key post‐release aspects of seed fate: seed destination (dispersed or nondispersed) and seed burial (buried or not buried). Experiments were performed in four different regions in the Iberian Peninsula. After three days of exposure of seeds to the main predator (fieldmice Apodemus sylvaticus), ca 30% of the seeds were removed. Seed destination affected the proportion of seeds escaping predation, but the sign, magnitude and statistical significance of the effect varied among the geographical regions. In the southern region (Cazorla), seeds dispersed in ant nests or intermediate destinations suffered scarcely any predation, but seeds under reproductive‐age plants experienced losses ca 50%. Conversely, in the northern region (Caurel), seeds in nests suffered significantly greater losses than seeds under plants or intermediate destinations, suggesting that nests were especially unsafe destinations. Seed burial had a strong impact on seed escape from predators, and its effect was highly consistent among geographical regions. In view of the consistency of its effect at different spatial scales, seed burial was a more general mechanism for predation avoidance than seed relocation to ant nests, which was habitat‐ and/or ant‐species‐dependent. Our results thus only partially support the predator‐avoidance hypothesis for the evolution of myrmecochory.