Impacts of the invasive Argentine ant on native ants and other invertebrates in coastal scrub in south‐eastern Australia

Invasive ants threaten native biodiversity and ecosystem function worldwide. Although their principal direct impact is usually the displacement of native ants, they may also affect other invertebrates. The Argentine ant, Linepithema humile (Dolichoderinae), one of the most widespread invasive ant species, has invaded native habitat where it abuts peri‐urban development in coastal Victoria in south‐eastern Australia. Here we infer impacts of the Argentine ant on native ants and other litter and ground‐dwelling invertebrates by comparing their abundance and taxonomic composition in coastal scrub forest either invaded or uninvaded by the Argentine ant. Species composition of native ants at bait stations and extracted from litter differed significantly between Argentine ant‐invaded and uninvaded sites and this was consistent across years. Argentine ants had a strong effect on epigeic ants, which were either displaced or reduced in abundance. The native ant Rhytidoponera victoriae (Ponerinae), numerically dominant at uninvaded sites, was completely absent from sites invaded by the Argentine ant. However, small hypogeic ants, including Solenopsis sp. (Myrmicinae) and Heteroponera imbellis (Heteroponerinae), were little affected. Linepithema humile had no detectable effect upon the abundance and richness of other litter invertebrates. However, invertebrate group composition differed significantly between invaded and uninvaded sites, owing to the varied response of several influential groups (e.g. Collembola and Acarina). Floristics, habitat structure and measured environmental factors did not differ significantly between sites either invaded or uninvaded by Argentine ants, supporting the contention that differences in native ant abundance and species composition are related to invasion. Changes in the native ant community wrought by Argentine ant invasion have important implications for invertebrate communities in southern Australia and may affect key processes, including seed dispersal.     

Effect of Argentine ant invasions on ground-dwelling arthropods in northern California riparian woodlands

Although the Argentine ant (Linepithema humile) is a widespread invasive species that displaces native ants throughout its introduced range, the effects of these invasions on arthropods other than ants remain poorly known. This study documents the consequences of Argentine ant invasions on ants and other ground-dwelling arthropods in northern California riparian woodlands. Baits and unbaited pitfall traps were used to sample different components of the arthropod communities at five pairs of uninvaded and invaded sites. Sites occupied by Argentine ants supported almost no native epigeic ants except for the winter-active Prenolepis imparis. Sites with Argentine ants averaged four to ten times more ant workers than did sites with native ants, but ant worker biomass did not differ between uninvaded and invaded sites. Argentine ants recruited to baits in invaded areas, on average, in less than half the time of native ants in uninvaded areas. Despite the loss of epigeic native ants, higher Argentine ant worker abundance, and faster recruitment by Argentine ants at invaded sites, pitfall trap samples from uninvaded and invaded areas contained similar abundances and diversities of non-ant arthropods. These findings suggest that Argentine ants and the native ants they displace interact with the ground-dwelling arthropods of these habitats in a similar manner. Received: 24 February 1997 / Accepted: 9 November 1997     

Impacts of Argentine ants on invertebrate communities with below-ground consequences

The Argentine ant, Linepithema humile (Mayr), is an invasive species that has been associated with various negative impacts in native communities around the world. These impacts, as for other invasive ants, are principally towards native ant species, and impacts on below-ground processes such as decomposition remain largely unexplored. We investigated the relationship between Argentine ants and invertebrate fauna, litter decomposition and soil microbial activity between paired invaded and uninvaded sites at two locations in Auckland, New Zealand, where there has been no research to date on their impacts. We examined the diversity and composition of invertebrate and microorganisms communities, and differences in soil and litter components. The composition of invertebrates (Order-level, ant and beetle species) was different between invaded and uninvaded sites, with fewer ants, isopods, amphipods, and fungus-feeding beetles at the invaded sites, whereas Collembola were more abundant at the invaded sites. There were significant differences in soil chemistry, including higher carbon and nitrogen microbial biomass at uninvaded sites. Several litter components were significantly different for Macropiper excelsum. The fibre content of litter was higher, and key nutrients (e.g. nitrogen) were lower, at invaded sites, indicating less breakdown of litter at invaded sites. A greater knowledge of the history of invasion at a site would clarify variation in the impacts of Argentine ants, but their persistence in the ground litter layer may have long-term implications for soil and plant health in native ecosystems.     

Invasion of Argentine ants (Hymenoptera: Formicidae) in South Australia: Impacts on community composition and abundance of invertebrates in urban parklands

Abstract The Argentine ant (Linepithema humile Mayr) is a worldwide invasive pest species that has been associated with losses of native ant and non‐ant invertebrates in its introduced range. To date, few studies have investigated the effects of Argentine ants on native invertebrates in Australia. This study assessed the effects of Argentine ants on community composition of invertebrates, with particular focus on resident ant communities and functional groups. In this study, the author compared the composition and abundances of invertebrates between invaded and uninvaded locations at four paired sites in Adelaide, South Australia. The results showed that there were significantly fewer non‐Argentine ants at invaded sites than at uninvaded sites. In particular, ants from the two common and widespread genera Iridomyrmex and Camponotus showed decreased abundances at the invaded sites. Multidimensional scaling analyses revealed differences in the composition of ant communities at the invaded and uninvaded sites, with uninvaded sites characterized by a similar native ant species composition, while communities at the invaded sites displayed much greater variability in species composition. These results suggest that the presence of Argentine ants may have a negative effect on particular ant genera and functional groups, with likely disruptions to ecosystem processes.     

Native predators living in invaded areas: responses of terrestrial amphibian species to an Argentine ant invasion

Predator–prey interactions play a key role in the success and impacts of invasive species. However, the effects of invasive preys on native predators have been poorly studied. Here, we first reviewed hypotheses describing potential relationships between native predators and invasive preys. Second, we examined how an invasive prey, the Argentine ant (Linepithema humile), affected a native terrestrial amphibian community. In the field, we looked at the structure of the amphibian community in invaded versus uninvaded areas and characterized amphibian trophic ecology. The amphibian community sampled seemed to show a species-dependent response in abundance to invasion: adults of the natterjack toad (Bufo calamita), the species demonstrating the highest degree of ant specialization, were less abundant in invaded areas. Although available ant biomass was significantly greater in invaded than in uninvaded areas (only Argentine ants occurred in the former), amphibians consumed relatively fewer ants in invaded areas. In the lab, we quantified amphibian consumption of Argentine ants versus native ants and assessed whether consumption patterns could have been influenced by prior exposure to the invader. The lab experiments corroborated the field results: amphibians preferred native ants over Argentine ants, and prior exposure did not influence consumption. Differences in preference explained why amphibians consumed fewer Argentine ants in spite of their greater relative availability; they might also explain why the most ant-specialized amphibians seemed to avoid invaded areas. Our results suggest the importance to account for predator feeding capacities and dietary ranges to understand the effects of invasive species at higher trophic levels.     

Consumption of introduced prey by native predators: Argentine ants and pit-building ant lions

When populations of native predators are subsidized by numerically dominant introduced species, the structure of food webs can be greatly altered. Surprisingly little is known, however, about the general factors that influence whether or not native predators consume introduced species. To learn more about this issue, we examined how native pit-building ant lions (Myrmeleon) are affected by Argentine ant (Linepithema humile) invasions in coastal southern California. Compared to areas without L. humile, invaded areas contained few native ant species and were deficient in medium-sized and large bodied native ants. Based on these differences, we predicted that Argentine ants would negatively affect ant lion larvae. Contrary to this expectation, observational surveys and laboratory growth rate experiments revealed that Myrmeleon were heavier, had longer mandibles, and grew more quickly when their main ant prey were Argentine ants rather than native ants. Moreover, a field transplant experiment indicated that growth rates and pupal weights were not statistically different for larval ant lions reared in invaded areas compared to those reared in uninvaded areas. Argentine ants were also highly susceptible to capture by larval Myrmeleon. The species-level traits that presumably make Argentine ant workers susceptible to capture by larval ant lions—small size and high activity levels—appear to be the same characteristics that make them unsuitable prey for vertebrate predators, such as horned lizards. These results underscore the difficulties in predicting whether or not numerically dominant introduced species serve as prey for native predators.     

Effects of Argentine ants (Hymenoptera: Formicidae) on myrmecophilous lycaenid butterfly,Narathura bazalus (Lepidoptera: Lycaenidae), in western Japan

Effects of the invasive Argentine ant, Linepithema humile (Mayr), on a myrmecophilous butterfly, Narathura bazalus (Hewiston), were investigated in the field in western Japan. Larvae of N. bazalus were attended by workers of Argentine ants in invaded parks and of ten native and one cosmopolitan ant species in uninvaded parks. The abundance of eggs and larvae were not significantly different between invaded and uninvaded parks. Pupal weight and parasitized ratio by tachinid flies were also not different between the two types of parks. These results indicate that the role of Argentine ants for the butterfly might be almost equivalent to the native ants.     

Scavenging in Mediterranean ecosystems: effect of the invasive Argentine ant

Above-ground invertebrates may represent a high proportion of animal biomass, but few data are available on their fate after death. In Mediterranean ant communities, they are frequently scavenged by ants. Here, we assessed the consequences of Argentine ant invasion on the removal of arthropod corpses in Doñana National Park (SW Spain). In three natural habitats that differed in their degree of vegetation cover (i.e. protection for ants against high temperatures), we experimentally provided dead Drosophila, and observed their disappearance over a 60-min period at different times of day and year. The habitats used were isolated cork oak trees, pine tree forest and dry scrubland; we compared invaded with uninvaded plots in each. Oak trees were the most invaded habitat, while scrubland was the least and the only one where the Argentine ant coexisted with native ant species. In accordance with this degree of invasion, the Argentine ant removed the highest percentage of dead flies in oak trees and the lowest in scrubland. Its performance as scavenger was higher than uninvaded ant communities, but it was reduced at high temperatures, when native species were highly efficient. The saturated distribution of the Argentine ant colony seems to be the key to its efficiency. We discuss how the occurrence and scavenger efficiency of the Argentine ant could affect the nutrient cycling and the progression of its invasion.     

Rainfall facilitates the spread, and time alters the impact, of the invasive Argentine ant

Climate change may exacerbate invasions by making conditions more favorable to introduced species relative to native species. Here we used data obtained during a long-term biannual survey of the distribution of ant species in a 481-ha preserve in northern California to assess the influence of interannual variation in rainfall on the spread of invasive Argentine ants, Linepithema humile, and the displacement of native ant species. Since the survey began in 1993, Argentine ants have expanded their range into 74 new hectares. Many invaded hectares were later abandoned, so the range of Argentine ants increased in some years and decreased in others. Rainfall predicted both range expansion and interannual changes in the distribution of Argentine ants: high rainfall, particularly in summer months, promoted their spread in the summer. This suggests that an increase in rainfall will promote a wider distribution of Argentine ants and increase their spread into new areas in California. Surprisingly, the distribution of two native ant species also increased following high rainfall, but only in areas of the preserve that were invaded by L. humile. Rainfall did not have a negative impact on total native ant species richness in invaded areas. Instead, native ant species richness in invaded areas increased significantly over the 13 years of observation. This suggests that the impact of Argentine ants on naïve ant communities may be most severe early in the invasion process.     

Factors governing rate of invasion: a natural experiment using Argentine ants

Predicting the success of biological invasions is a major goal of invasion biology. Determining the causes of invasions, however, can be difficult, owing to the complexity and spatio-temporal heterogeneity of the invasion process. The purpose of this study was to assess factors influencing rate of invasion for the Argentine ant (Linepithema humile), a widespread invasive species. The rate of invasion for 20 independent Argentine ant populations was measured over 4 years in riparian woodlands in the lower Sacramento River Valley of northern California. A priori predictors of rate of invasion included stream flow (a measure of abiotic suitability), disturbance, and native ant richness. In addition, baits were used to estimate the abundance of Argentine ants and native ants at the 20 sites. A multiple regression model accounted for nearly half of the variation in mean rate of invasion (R ² = 0.46), but stream flow was the only significant factor in this analysis. Argentine ants spread, on average, 16 m year⁻¹ at sites with permanent stream flow and retreated, on average, −6 m year⁻¹ at sites with intermittent stream flow. Rate of invasion was independent of both disturbance and native ant richness. Argentine ants recruited to more baits in higher numbers in invaded areas than did native ants in uninvaded areas. In addition, rate of invasion was positively correlated with the proportion of baits recruited to by native ants in uninvaded areas. Together, these findings suggest that abiotic suitability is of paramount importance in determining rate of invasion for the Argentine ant. Received: 16 September 1997 / Accepted: 8 February 1998     

Seed removal decrease by invasive Argentine ants in a high Nature Value farmland

Seed dispersal by ants is an important ecological process that maintains the structure anddiversity of natural communities, however, it is vulnerable to biological invasions. Argentine ants are one of the worst invasive ant species and cause severe changes in ecosystem processes and native ant biodiversity declines in invaded sites. Here, we studied seed removal by ants combining observations and a cafeteria experiment with seeds of four myrmecochorous plant species (Centaurea sphaerocephala, Rosmarinus officinalis, Silybum marianum, and Ulex australis) in two sites (invaded and uninvaded) located in the Mediterranean Montado ecosystem and classified as High Nature Value farmland (HNV). Significant differences in daily seed removal rates were found between the two study sites. In uninvaded sites, several native ant species were attracted to the seeds, resulting in all seeds being removed rapidly. The majority of seed removal events were carried out by two key seed disperses Pheidole pallidula (71%) and Aphaenogaster iberica (26%) with a clear preference for diaspored with larger and heavier elaiosome (i.e., C. sphaerocephala, S. marianum). By contrast, while the Argentine ant showed some interest (68% of seeds were interacted with), no seed removal events were observed. The extirpation of the local ant fauna by the Argentine ant and its inability to ensure seed dispersal services may lead to the interference and eventually to the collapse of seed dispersal of the four studied myrmecochorous plants in the invaded site in the future. We argue that these discrete but severe consequences of an invasive species on a key ecological process may strongly affect the functioning of the Montado ecosystem.     

The Argentine ant persists through unfavorable winters via a mutualism facilitated by a native tree

Mutualisms and facilitations can fundamentally change the relationship between an organism's realized and fundamental niche. Invasive species may prove particularly suitable models for investigating this relationship as many are dependent on finding new partners for successful establishment. We conducted field-based experiments testing whether a native tree facilitates the successful survival of the invasive Argentine ant, Linepithema humile (Mayr), through unfavorable winter conditions in the southeastern United States. We found Argentine ant nests aggregated around the native loblolly pine, Pinus taeda L., during the winter months. The bark of this tree absorbed enough radiant solar energy to reach temperatures suitable for Argentine ant foraging even when ambient temperatures should have curtailed all foraging. Conversely, foraging ceased when the trunk was shaded. The sun-warmed bark of this tree gave the Argentine ant access to a stable honeydew resource. Argentine ants were not found on or near deciduous trees even though bark temperatures were warm enough to permit Argentine ant foraging on cold winter days. Augmenting deciduous trees with sucrose water through the winter months lead to Argentine ant nests remaining at their base and Argentine ants foraging on the tree. The Argentine ant requires both foraging opportunity and a reliable winter food source to survive through unfavorable winter conditions in the southeastern United States. The loblolly pine provided both of these requirements extending the realized niche of Argentine ants beyond its fundamental niche.     

Infiltration of a facultative ant–plant mutualism by the introduced Argentine ant: effects on mutualist diversity and mutualism benefits

1. Ant–plant mutualisms have been the focus of considerable empirical research, but few studies have investigated how introduced ants affect these interactions. Using 2 years of survey data, this study examines how the introduced Argentine ant [Linepithema humile (Mayr)] differs from native ants with respect to its ability to protect the extrafloral nectary‐bearing coast barrel cactus (Ferocactus viridescens) in Southern California. 2. Eighteen native ant species visited cacti in uninvaded areas, but cacti in invaded areas were primarily visited by the Argentine ant. The main herbivore of the coast barrel cactus present at the study sites is a leaf‐footed bug (Narnia wilsoni). 3. Herbivore presence (the fraction of surveys in which leaf‐footed bugs were present on individual cacti) was negatively related to ant presence (the fraction of surveys in which ants were present on individual cacti). Compared with cacti in uninvaded areas, those in invaded areas were less likely to have herbivores and when they did had them less often. 4. Seed mass was negatively related to herbivore presence, and this relationship did not differ for cacti in invaded areas versus those in uninvaded areas. 5. Although the Argentine ant might provide superior protection from herbivores, invasion‐induced reductions in ant mutualist diversity could potentially compromise plant reproduction. The cumulative number of ant species on individual cacti over time was lower in invaded areas and was associated with a shortened seasonal duration of ant protection and reduced seed mass. These results support the hypothesis that multiple partners may enhance mutualism benefits.     

Abiotic factors control invasion by Argentine ants at the community scale

1. A prominent and unresolved question in ecology concerns why communities differ in their susceptibility to invasion. While studies often emphasize biotic resistance, it is less widely appreciated how the physical environment affects community vulnerability to invasion. 2. In this study we performed field experiments to test how abiotic variation directly and indirectly influences the extent to which Linepithema humile Mayr (Argentine ants) invade seasonally dry environments in southern California. 3. In controlled and replicated experiments involving drip irrigation, we demonstrate (i) that elevated levels of soil moisture increased both the abundance of Argentine ants and their ability to invade native ant communities and (ii) that cessation of irrigation caused declines in the abundance of Argentine ants and led to their withdrawal from previously occupied areas. 4. Because drip irrigation stimulated plant growth, in an additional experiment we manipulated both soil moisture and plant cover to assess the direct vs. indirect effects of added water on the abundance of L. humile. 5. Local abundance of Argentine ants increased in irrigated plots but was 38% higher in irrigated plots with plants compared to irrigated plots where plant growth was suppressed. The results of this experiment thus argue for a direct role of soil moisture in influencing Argentine ant abundance but suggest that that the indirect effects of added water may also be important. 6. Our study illustrates more generally that fine-scale variation in the physical environment can control whether communities become invaded by non-native species and suggests that an understanding of community susceptibility to invasion will be improved by a better appreciation of interactions between the biotic and abiotic environment.     

Introduced ants reduce interaction diversity in a multi‐species,ant–aphid mutualism

Mutualisms contribute in fundamental ways to the origin, maintenance and organization of biological diversity. Introduced species commonly participate in mutualisms, but how this phenomenon affects patterns of interactions among native mutualists remains incompletely understood. Here we examine how networks of interactions among aphid‐tending ants, ant‐tended aphids, and aphid‐attacking parasitoid wasps differ between 12 spatially paired riparian study sites with and without the introduced Argentine ant Linepithema humile in southern California. To resolve challenges in species identification, we used DNA barcoding to identify aphids and screen for parasitoid wasps (developing inside their aphid hosts) from 170 aphid aggregations sampled on arroyo willow Salix lasiolepis. Compared to uninvaded sites, invaded sites supported significantly fewer species of aphid‐tending ants and ant‐tended aphids. At invaded sites, for example, we found only two species of ant‐tended aphids, which were exclusively tended by L. humile, whereas at uninvaded sites we found 20 unique ant–aphid interactions involving eight species of ant‐tended aphids and nine species of aphid‐tending ants. Ant–aphid linkage density was thus significantly lower at invaded sites compared to uninvaded sites. We detected aphid parasitoids in 14% (28/198) of all aphid aggregations. Although the level of parasitism did not differ between invaded and uninvaded sites, more species of wasps were detected within uninvaded sites compared to invaded sites. These results provide a striking example of how the assimilation of introduced species into multi‐species mutualisms can reduce interaction diversity with potential consequences for species persistence.     

Impacts of Argentine ants on avian nesting success

Biological invasions can have severe and widespread impacts on ecological communities. A few species of ants have become particularly damaging invaders but quantitative data of their impacts on many taxa is still lacking. We provide experimental evidence using artificial nests baited with quail eggs that the invasive Argentine ant (Linepithema humile) can be a significant avian nest predator – Argentine ants recruited to more nests and in higher abundance than the native ant species they displace. However, at a site invaded by Argentine ants, we monitored over 400 nests of a ground-nesting species, the Dark-eyed Junco (Junco hyemalis), and found that less than 2% of nests failed as a result of Argentine ant predation/infestation. A review of the literature also suggests that Argentine ants may not be a serious threat to bird nests relative to other predators or parasites. However, invasive ants with the capability of overwhelming prey though stinging (specifically the red-imported fire ant, Solenopsis invicta), may have a higher impact on avian nesting success. Received 14 January 2005; revised 28 April 2005; accepted 12 May 2005.     

Behavioral interactions of the invasive Argentine ant with native ant species

Summary: The Argentine ant, Linepithema humile, has invaded many areas of the world, displacing native ants. Its behavior may contribute to its competitive success. Staged and natural encounters were observed at food resources in the field, between Argentine ants and eight ant species native to northern California. There was no relation between the frequency of aggression by any ant species and the outcome of encounters, though Argentine ants were more likely than ants of native species to behave aggressively. When an ant of one species initiated an encounter of any kind with an ant of another species, the ant that did not initiate was likely to retreat. This was true of all species studied. Most encounters between ants were initiated by Argentine ants. Thus the native species tended to retreat more frequently than Argentine ants. Interactions between Argentine ants and native species at food resources, causing ants of native species to retreat, may help Argentine ants to displace native species from invaded areas.     

Exploitation and interference competition between the invasive Argentine ant,Linepithema humile,and native ant species

Interactions between the invasive Argentine ant, Linepithema humile, and native ant species were studied in a 450-ha biological reserve in northern California. Along the edges of the invasion, the presence of Argentine ants significantly reduced the foraging success of native ant species, and vice versa. Argentine ants were consistently better than native ants at exploiting food sources: Argentine ants found and recruited to bait more consistently and in higher numbers than native ant species, and they foraged for longer periods throughout the day. Native ants and Argentine ants frequently fought when they recruited to the same bait, and native ant species were displaced from bait during 60% of these encounters. In introduction experiments, Argentine ants interfered with the foraging of native ant species, and prevented the establishment of new colonies of native ant species by preying upon winged native ant queens. The Argentine ants' range within the preserve expanded by 12 ha between May 1993 and May 1994, and 13 between September 1993 and September 1994, with a corresponding reduction of the range of native ant species. Although some native ants persist locally at the edges of the invasion of Argentine ants, most eventually disappear from invaded areas. Both interference and exploitation competition appear to be important in the displacement of native ant species from areas invaded by Argentine ants.     

Sampling across space and time to validate natural experiments: an example with ant invasions in Hawaii

Measuring the impacts of invasive species on recipient biotic communities can be challenging because standard ecological field experiments may be impossible, undesirable or inaccurate due to limitations of scale. Many studies therefore rely on a comparative observational approach, in which invaded areas are compared to similar nearby uninvaded areas, but this approach suffers from reduced inferential power because of the inability to randomly assign treatments. When possible, the best solution may be to incorporate comparative sampling over time, from before and after invasion, as a means of validating comparisons across space. We used this approach to assess the impacts of Argentine and big-headed ants on arthropod communities at two natural area sites in the Hawaiian Islands. Two sampling events, separated by 2 years, indicated that invaded plots at each site were significantly different from uninvaded plots in terms of overall community composition and distributions of certain indicator species. This compositional change associated with ant invasion was accompanied by a significant decrease in arthropod species richness at one site, but not at the second site. Changes in arthropod communities from before invasion to after invasion in separate before–after plots at each site were qualitatively consistent with the aforementioned changes measured across the invasion boundaries, but were smaller in magnitude because the invasion processes were still in early stages. Together, these findings support the inference that carefully planned studies of biological invasions that use a comparative observational approach can provide an accurate picture of resultant community level changes, and illustrate an effective method for validating such studies.     

Invasive Argentine ants (<Emphasis Type="Italic">Linepithema humile</Emphasis>) do not replace native ants as seed dispersers of <Emphasis Type="Italic">Dendromecon rigida</Emphasis> (Papaveraceae) in California,USA

We investigated the indirect effects of Argentine ant (Linepithema humile) invasions on patterns of seed dispersal and predation in the myrmecochorous tree poppy Dendromecon rigida in coastal San Diego County, California. Significantly more seeds were removed from ant-accessible seed stations at sites numerically dominated by a common harvester ant (Pogonomyrmex subnitidus), a native disperser of these seeds and a species sensitive to displacement by L. humile, than from those stations at sites where L. humile was in the majority. Predation of seeds was high, but variable, across sites, suggesting that reduced dispersal could result in increased seed predation in some habitats. Removal of elaiosomes did not affect the frequency with which predators removed seeds, but ants removed significantly more seeds with elaiosomes than without. In behavior trials, only P. subnitidus was able to carry seeds of Dendromecon rigida effectively. L. humile and a small native ant species, Dorymyrmex insanus, while displaying interest in the diaspores, were seldom able to carry whole seeds and, when they did, only carried them a few centimeters. Displacement of native harvester ants by L. humile appears to decrease the dispersal of Dendromecon rigida seeds and may be increasing loss of seeds due to predation.