Seed viability and germination success of Acacia tortilis along land‐use and aridity gradients in the Eastern Sahara

Our study focuses on the keystone species Acacia tortilis and is the first to investigate the effect of domestic ungulates and aridity on seed viability and germination over an extensive part of the Eastern Sahara. Bruchids infest its seeds and reduce their viability and germination, but ingestion by ruminant herbivores diminishes infestation levels and enhances/promotes seed viability and germination. The degree of these effects seems to be correlated with animal body mass. Significantly reduced numbers of wild ruminant ungulates have increased the potential importance of domestic animals and pastoral nomadism for the functionality of arid North African and Middle Eastern ecosystems. We sampled seeds (16,543) from A. tortilis in eight areas in three regions with different aridity and land use. We tested the effect of geography and sampling context on seed infestation using random effects logistic regressions. We did a randomized and balanced germination experiment including 1193 seeds, treated with different manure. Germination time and rates across geography, sampling context, and infestation status were analyzed using time‐to‐event analyses, Kaplan–Meier curves and proportional hazards Cox regressions. Bruchid infestation is very high (80%), and the effects of context are significant. Neither partial infestation nor adding manure had a positive effect on germination. There is a strong indication that intact, uningested seeds from acacia populations in the extremely arid Western Desert germinate more slowly and have a higher fraction of hard seeds than in the Eastern Desert and the Red Sea Hills. For ingested seeds in the pastoralist areas we find that intact seeds from goat dung germinate significantly better than those from camel dung. This is contrary to the expected body‐mass effect. There is no effect of site or variation in tribal management.     

Effect of parasitoids,seed‐predators and ant‐mutualists on fruiting success and germination of Acacia drepanolobium in Kenya

Acacia drepanolobium is an obligate ant‐plant that bears dehiscent pods exploited by predispersal seed‐predators and parasitoids. Fruit set and seed germination in relation to ant‐association, bruchid and parasitoid infestation were investigated in a large‐scale multi‐year study in Kenya. Ant‐association had a significant impact on the overall numbers of fruiting trees. 94.8% of Crematogaster mimosae‐occupied trees, 25.6% of C. nigriceps‐occupied trees, and 82.2% of Tetraponera penzigi‐occupied trees set fruit. Within each tree, ant‐association had no significant impact on the amount of seed produced: C. mimosae‐occupied trees produced 47.5 seeds per branch, C. nigriceps‐occupied trees 44.9 seeds per branch and T. penzigi‐occupied trees 38.3 seeds per branch. Ant‐association did not limit seed‐predators or vary significantly by ant. Seeds infested by bruchids germinated in significantly lower proportions (6.2%) compared to uninfested seed (78.6%). Bruchid‐infested seed is also exploited by parasitoid wasps. Parasitoids appear to have a moderate but significant ‘rescue’ effect on bruchid‐infested seed with 18.4% of parasitoid‐infested seed germinating. Stable isotopes (δ15N) revealed the trophic structure of the seed‐associated insects, showing clearly that bruchids are seed predators and the parasitoid wasp Dinarmus magnus exploits the dominant Bruchidius sp.     

Fragmentation and Alteration of Seed Dispersal Processes: An Initial Evaluation of Dung Beetles, Seed Fate, and Seedling Diversity1

Given current accelerated trends of tropical land conversion, forest fragments are being incorporated into many conservation programs. For investing in fragments to be a viable conservation strategy, forest fragments must maintain their ecological integrity over the long term. Based on fieldwork in 22 forest fragments in the crater lakes region of western Uganda and in the continuous forest of Kibale National Park, we examined (1) seed predation on experimentally dispersed seeds, (2) abundance and composition of the dung beetle community that may play a major role in removing seeds from sites of high seed predation, and (3) compared the fragments’ seedling community composition to adult tree community composition and the seedling community in continuous forest. First, the rate of seed removal at experimental stations was lower in forest fragments (85% remaining after 1 day) than at stations in the continuous forest (79% remaining) and the probability of stations being discovered by seed predators was lower in fragments (23%) than in the intact forest (41%). Second, there was a 62 percent decline in fragment dung beetle abundance. The magnitude of this decline varied among dung beetle guilds that process dung and seeds in different fashions. The abundance of large rollers that move large seeds away from sites of defecation did not differ, but medium and smaller rollers and burying beetles that process small and medium‐sized seeds were less common in the fragments than in the intact forest. Finally, we compared the seedling community composition relative to adult tree community composition by identifying all adult trees in each fragment and by sampling the composition of the seedling community. We found some evidence to suggest that there was movement of seeds among forest fragments by large‐bodied dispersers, particularly chimpanzees (Pan troglodytes) and hornbills (Ceratogymna subcylindricus).     

Large African herbivores,bruchid beetles and their interactions with Acacia seeds

This study investigated the interactions of large African herbivores and bruchid seed beetles with Acacia seeds. The germination of bruchid-infested and uninfested seeds was compaed. The effects of pod consumption by large herbivores on bruchid infestation and seed germination was also assessed. Bruchid-infested seeds did germinate, and the germination of bruchid-infested and uninfested A. tortilis, A. nilotica and A. hebeclade seeds did not differ. Pod ingestion by large herbivores lowered the bruchid infestation of consumed and defaecated seeds compared to uningested seeds. Uninfested, ingested and voided A. tortilis seeds germinated seeds. Furthermore, infested A. tortilis seeds egested by giraffe, kudu and ostrich germinated better than infested, uningested seeds. Pod ingestion by large herbivores may reduce bruchid infestation, increase Acacia seed germination and therefore increase potential Acacia seedling recruitment.     

Maternal effects on the life histories of bruchid beetles infesting Acacia raddiana in the Negev desert, Israel

Acacia trees (Leguminosae) in the Negev desert of Israel are in substantial danger of extinction as a result of high mortality, caused by anthropogenically induced water stress and very low recruitment, which is highly negatively affected by bruchid beetle infestation. Intensive seed infestation (up to 97%) by bruchids on Acacia species in the Negev desert has been described. We hypothesized that water‐stressed trees would be less able to produce secondary defense compounds in their seeds to decrease seed herbivory and hence suffer higher infestation. Thus, there should be a negative correlation between the beetle's fitness and the tree's physiological state. We further hypothesized the mechanism of the correlation to be maternal effects on beetle quality. Thus, beetles whose mothers were reared on seeds of trees in a poor state should have higher fitness, regardless of the quality of the seeds in which they were reared. We reared F₁ generation Caryedon palaestinicus Southgate (Coleoptera: Bruchidae) in seeds of Acacia raddiana Savi (Leguminosae) trees in good and in poor physiological state. We then conducted a reciprocal infestation experiment enabling females to lay on seeds from both sources. We found maternal effects on offspring number but not on body mass. The major effect was the limited survival of maternal beetles on trees in good physiological state. However, contrary to our prediction, C. palaestinicus developing in seeds from Acacia raddiana in good physiological state had significantly higher body mass.     

Telemetric tracking of scatterhoarding and seed fate in a Central African forest

In seed predation studies, removal of a seed is only the first step of a dynamic process that may result in dispersal rather than seed death. This process, termed seed fate, has received little attention in African forests, particularly in Central Africa. We experimentally assessed the initial steps of seed fate for two tree species—the large‐seeded Pentaclethra macrophylla and the relatively small‐seeded Gambeya lacourtiana—in northeastern Gabon. Specifically, we evaluated whether seed size and seed consumer identity are important determinants of seed fate. We established experimental stations under conspecific fruiting trees, each comprising three seeds fitted with telemetric thread tags to facilitate their recovery, and a motion‐sensitive camera to identify visiting mammals. In total, animals removed 76 tagged seeds from experimental stations. Small Murid rats and mice primarily removed small Gambeya seeds, whereas large‐bodied rodents and mandrills primarily removed large Pentaclethra seeds. Gambeya seeds were carried shorter distances than Pentaclethra seeds and were less likely to be cached. The two large‐bodied rodents handled seeds differently: Cricetomys emini larderhoarded nearly all (N = 15 of 16) encountered Pentaclethra seeds deep in burrows, while Atherurus africanus cached all (N = 5 of 5) encountered Pentaclethra seeds singly under 1–3 cm of leaf litter and soil, at an average distance of 24.2 m and a maximum distance of 46.3 m from experimental stations. This study supports the hypothesis that seed fate varies based on seed size and seed consumer identity, and represents the first telemetric experimental evidence of larderhoarding and scatterhoarding in the region.     

The production, storage and viability of seeds of Acacia karroo and A. nilotica in a grassy savanna in KwaZulu-Natal, South Africa

African Acacias are often major contributors to the progressive increase in the woody component of savannas, a phenomenon commonly referred to as bush encroachment. They produce large quantities of seed and may have large soil‐stored seed banks. In Hluhluwe–Umfolozi Park, the number of adult Acacia nilotica trees per hectare far exceed that of A. karroo adults. The relative dominance is reversed in the juvenile stage with A. karroo outnumbering A. nilotica threefold outside closed woodlands. Acacia karroo trees were smaller than A. nilotica trees on average, but produced more seeds for a given basal diameter size class. Acacia karroo showed less bruchid infestation than A. nilotica at all stages of pod development. Unlike A. nilotica, a proportion of A. karroo seeds was able to germinate after bruchid attack. We detected no difference between the two species in the soil‐stored seed bank or in the viability of seeds found in the seed bank.     

Tamarins and Dung Beetles: An Efficient Diplochorous Dispersal System in the Peruvian Amazonia

Dung beetles fulfill several key functions in ecosystems but their role as secondary seed dispersers is probably one of the most complex ones. Various factors, such as seed characteristics, dispersal pattern induced by the primary disperser, season, and habitat, can affect the seed–beetle interaction. Particularly little is known about the fate of seeds primarily dispersed in small feces. The aim of this study was to investigate the effects of these factors on the dung beetle community (species composition, number and size of individuals) and its consequences on burial occurrence and depth of seeds primarily dispersed by two tamarin species. We captured dung beetles in a Peruvian rain forest with 299 dung‐baited pitfall traps to characterize the dung beetle community. Seed burial occurrence and depth were assessed by marking in situ 551 dispersed seeds in feces placed in cages. Among these seeds, 22.5 percent were buried by dung beetles after 2 d. We observed a significant effect of the amount of dung, season, time of deposition, and habitat on the number of individuals and species of dung beetles, as well as on seed burial occurrence and depth, while the tamarin species significantly influenced only the number and the size of dung beetles. This seed dispersal loop is particularly important for forest regeneration: small to large seeds dispersed by tamarins in secondary forest can be buried by dung beetles. These seeds can thus benefit from a better protection against predation and a more suitable microenvironment for germination, potentially enhancing seedling recruitment.     

Frugivory and Seed Dispersal by Brown Lemurs in a Malagasy Tropical Dry Forest

In the Ankarafantsika tropical dry forest (northwestern Madagascar), the common brown lemur (Eulemur fulvus fulvus) is the largest frugivore and probably the sole disperser of large‐seeded plants (seed diameter > 10 mm). To investigate seed dispersal by this primate, I recorded the feeding activities of a troop; also conducted fecal analyses, germination trials on defecated seeds, and a vegetation survey over 1 yr (beginning Dec 2006). Brown lemurs mostly consumed fruit (68%). The fruit of Vitex beraviensis was the most exploited resource (21% of total feeding time). Among dung samples, 1126 contained intact seeds of 70 plant species, with a median of six seeds and two species per sample. These data indicate that the brown lemur population dispersed approximately 9854 seeds/km²/d. Although the number of annually defecated seeds was overwhelmingly the largest in Grewia triflora, many of the small seeds were often clumped in dung piles. In contrast, large seeds of V. beraviensis occurred in the largest number of dung samples. The rate and time of seed germination in V. beraviensis were improved by passage through brown lemur guts. Therefore, V. beraviensis may readily establish seedlings in sites of brown lemur fecal deposition. Vitex beraviensis and brown lemurs are probably involved in a strong mutualism. Twenty‐three large‐seeded plants were probably dependent on brown lemurs for seed dispersal and some of these species were common trees in the forest. Maintenance of these key plant–animal interactions will probably contribute to the conservation of species diversity and intact regeneration of the Ankarafantsika forest.     

Toucans (Ramphastos ambiguus) facilitate resilience against seed dispersal limitation to a large‐seeded tree (Virola surinamensis) in a human‐modified landscape

Large‐seeded plants may suffer seed dispersal limitation in human‐modified landscapes if seed dispersers are absent or unable to disperse their seeds. We investigated dispersal limitation for the large‐seeded tree Virola surinamensis in a human‐modified landscape in southern Costa Rica. During two fruiting seasons, we monitored crop size, seed removal rates, the number of fruiting conspecifics within 100 m, and feeding visitation rates by frugivores at trees located in high and low forest disturbance conditions. Seed removal rates and the total number of seeds removed were high regardless of the disturbance level, but these parameters increased with tree crop size and decreased with the number of fruiting V. surinamensis trees within a 100 m radius. Trees at low disturbance levels were more likely to be visited by seed dispersers. Black mandibled toucans (Ramphastos ambiguus) and spider monkeys (Ateles geoffroyi) were the most important seed dispersers, based on visitation patterns and seed removal rates. Spider monkey feeding visits were more frequent at high disturbance levels, but the monkeys preferentially visited isolated trees with large yields and surrounded by a low number of fruiting Virola trees within 100 m. Toucan visitation patterns were not constrained by any of the predictors and they visited trees equally across the landscape. We suggest that isolated and highly fecund Virola trees are an important food resource for spider monkeys in human‐modified landscapes and that toucans can provide resilience against seed dispersal limitations for large‐seeded plants in human‐modified landscapes in the absence of hunting.     

Post‐Dispersal Seed Removal and Germination Selected Tree Species Dispersed by Cebus capucinus on Barro Colorado Island,Panama1

Dispersal quality, an important component of seed disperser effectiveness, may strongly affect the rate of plant recruitment. Here we evaluated the quality of Cebus monkey dispersal by comparing the secondary removal fate and germination of fresh and Cebus‐ingested seeds of nine tree species on Barro Colorado Island, Panama. Overall, rates of secondary seed removal by vertebrates were low, with most Cebus defecations remaining undisturbed for extended periods on the forest floor. Only four of 30 feces were completely buried by dung beetles, and we found significantly higher vertebrate removal of defecated seeds than control seeds for only one species, Cordia bicolor. Seed germination varied greatly between plant taxa. Seeds of 3 out of 9 species showed significantly higher percent germination after monkey gut passage than control fresh seeds. Germination times tended to be shorter for defecated than for control seeds but were significantly different only for one of nine species, Cecropia insignis. Low rates of seed removal from Cebus feces, coupled with high germination probabilities, suggest high dispersal effectiveness for Cebus and contrasts strongly with patterns of post‐dispersal seed fate recorded for other primate species.     

Dispersal and predation of diaspores of Coula edulis Baill. in an evergreen forest of Gabon

The African walnut (Coula edulis) is a tree species of African evergreen forests, the seeds of which are collected and traded by African people. Many animal species consume African walnut diaspores; however, their roles as dispersers or predators have yet to be clarified. In this study, we present observations conducted in two different habitats of a Gabonese region over a 3‐year period. The originality of this research resides in the combination of three complementary approaches: (i) the use of camera‐traps (ii) the exploration of land rodent burrows and (iii) the examination of elephant dung. In total, 408 camera‐trap photographs have shown seven animal species involved in the dispersal/predation of C. edulis. Among these seven frugivorous species, the bush pig was found to be the main consumer and predator of seeds. Land rodents (Muridae) are potential predators, as they damaged the seeds and buried them deep in the soil. They may also play a role in the regeneration process as a result of the loss of seeds during transportation. Finally, no seeds appeared to emerge intact from elephant faeces. These results indicate that the natural regeneration rate of this tree species is low, unless other complex mechanisms are involved.     

Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds1

The effectiveness of a seed disperser depends on the quantity and quality of dispersal. The quality of dispersal depends in large part on factors that affect the post–dispersal fate of seeds, and yet this aspect of dispersal quality is rarely assessed. In the particular case of seed dispersal through endozoochory, the defecation pattern produced has the potential of affecting the fate of dispersed seeds and consequently, dispersal quality and effectiveness. In this study, I assessed the effects of dung presence and dung/seed densities on seed predation by rodents and secondary dispersal by dung beetles. In particular, I compared seed fates in clumped defecation patterns, as those produced by howler monkeys, with seed fates in scattered defecation patterns, as those produced by other frugivores. I also determined the prevalence of red howler monkeys (Alouatta seniculus) as seed dispersers at the plant community level in Central Amazonia by determining the number of species they dispersed in a 25–month period. I found that dung presence and amount affected rodent and dung beetle behavior. Seed predation rates were higher when dung was present, and when it was in higher densities. The same number of seeds was buried by dung beedes, in dumped versus scattered defecation patterns, but more seeds were buried when they were inside large dung–piles versus small piles. Seed density had no effect on rodent or dung beetle behavior. Results indicate that caution should be taken when categorizing an animal as a high or low quality seed disperser before carefully examining the factors that affect the fate of dispersed seeds. Red howler monkeys dispersed the seeds of 137 species during the study period, which is the highest yet reported number for an Alouatta species, and should thus be considered highly prevalent seed dispersers at the plant community level in Central Amazonian terra firme rain forests.     

Persistence of the effect of frugivore identity on post‐dispersal seed fate: consequences for the assessment of functional redundancy

Large frugivores play an important role as seed dispersers and their extinction may affect plant regeneration. The consequences of such extinctions depend on the likelihood of other species being functionally redundant and on how post‐dispersal events are affected. We assess the functional redundancy of two seed dispersers of the Atlantic Forest, the muriqui (Brachyteles arachnoides) and the tapir (Tapirus terrestris) through the comparison of their seed dispersal quality, taking into account post‐dispersal events. We compare tapirs and muriquis for: (1) the dung beetle community associated with their feces; (2) the seed burial probability and burial depth by dung beetles; and (3) the seed mortality due to predators or other causes according to burial depth. We determine how seed burial affects seed dispersal effectiveness (SDE) and compare the dispersal quality of four plant species dispersed by these frugivores. Muriqui feces attract 16‐fold more dung beetles per gram of fecal matter and seeds experience 10.5‐fold more burial than seeds in tapir feces. In both feces types, seed mortality due to predation decreases with burial depth but seed mortality due to other causes increases. Total seed mortality differ within plant species according to the primary disperser. Therefore, the effect of seed burial on SDE varies according to the plant species, burial depth, and primary disperser. As tapirs and muriquis differently affect the seed fate, they are not functionally redundant. Since the effect of the primary disperser persists into the post‐dispersal events, we should consider the cascading effects of these processes when assessing functional redundancy.     

Seed weight patterns of Acacia tortilis from seven seed provenances across Botswana

Laboratory‐based seed storage systems have been developed as an alternative to in situ conservation for indigenous woody plant species. However, interactions between seed quality and environmental variables must be known for each species prior to seed collection, storage and sowing to ensure effective conservation. This study investigated Acacia tortilis seed weight/quality patterns across seven Botswana seed provenances in relation to: soil nutrient status, altitude; latitude; slope angle; % grass cover; height and density of other woody plants nearby. The higher rainfall and relatively eutrophic seed provenances of north‐western Botswana (Chobe, Okavango and Makgadikgadi) were associated with large A. tortilis species and seeds, as well as higher densities of woody plants. Spatial variation in seed weights of A. tortilis was principally a function of rainfall and soil organic carbon. Although more work is required to establish the relationship between seed weight and germination rates for A. tortilis, this research suggests that seed collection should focus on sites with high levels of rainfall and soil organic carbon.     

Long‐distance pollen and seed dispersal and inbreeding depression in Hymenaea stigonocarpa (Fabaceae: Caesalpinioideae) in the Brazilian savannah

Hymenaea stigonocarpa is a neotropical tree that is economically important due to its high‐quality wood; however, because it has been exploited extensively, it is currently considered threatened. Microsatellite loci were used to investigate the pollen and seed dispersal, mating patterns, spatial genetic structure (SGS), genetic diversity, and inbreeding depression in H. stigonocarpa adults, juveniles, and open‐pollinated seeds, which were sampled from isolated trees in a pasture and trees within a forest fragment in the Brazilian savannah. We found that the species presented a mixed mating system, with population and individual variations in the outcrossing rate (0.53–1.0). The studied populations were not genetically isolated due to pollen and seed flow between the studied populations and between the populations and individuals located outside of the study area. Pollen and seed dispersal occurred over long distances (>8 km); however, the dispersal patterns were isolated by distance, with a high frequency of mating occurring between near‐neighbor trees and seeds dispersed near the parent trees. The correlated mating for individual seed trees was higher within than among fruits, indicating that fruits present a high proportion of full‐sibs. Genetic diversity and SGS were similar among the populations, but offspring showed evidence of inbreeding, mainly originating from mating among related trees, which suggests inbreeding depression between the seed and adult stages. Selfing resulted in a higher inbreeding depression than mating among relatives, as assessed through survival and height. As the populations are not genetically isolated, both are important targets for in situ conservation to maintain their genetic diversity; for ex situ conservation, seeds can be collected from at least 78 trees in both populations separated by at least 250 m.     

Role of small rodents in the seed dispersal process: Microcavia australis consuming Prosopis flexuosa fruits

Understanding the functional role of animal species in seed dispersal is central to determining how biotic interactions could be affected by anthropogenic drivers. In the Monte Desert, mammals play different functional roles in Prosopis flexuosa seed dispersal, acting as opportunistic frugivores (endozoochorous medium‐sized and large mammals) or seed hoarders (some small sigmodontine rodents). Our objective was assessing the functional role of Microcavia australis, a small hystricognathi rodent, in the fruit removal and seed deposition stages of P. flexuosa seed dispersal, compared to sympatric sigmodontine rodents. In situ, we quantified fruit removal by small rodents during non‐fruiting and fruiting periods, and determined the distance seeds were transported, particularly by M. australis. In laboratory experiments, we analysed how M. australis stores seeds (through scatter‐ or larder‐hoarding) and how many seeds are left in caches as living seeds, relative to previous data on sigmodontine rodents. To conduct field studies, we established sampling stations under randomly chosen P. flexuosa trees at the Ñacuñán Man and Biosphere Reserve. We analysed fruit removal by small rodents and seed dispersal distance by M. australis using camera traps focused on P. flexuosa fruits covered with wire screen, which only allowed entry of small animals. In laboratory trials, we provided animals with a known number of fruits and assessed seed conditions after removal. Small rodents removed 75.7% of fruit supplied during the non‐fruiting period and 53.2% during the fruiting period. Microcavia australis and Graomys griseoflavus were the main fruit removers. Microcavia australis transported seeds to a mean distance of 462 cm and cached seeds mainly in scatter‐hoards, similarly as Eligmodontia typus. All transported seeds were left in fruit segments or covered only by the endocarp, never as predated seeds. Microcavia australis disperses P. flexuosa seeds by carrying fruits away from a source to consume them and then by scatter‐hoarding fruits and seeds.     

Spatial spread and infestation risk assessment in the Asian longhorned beetle,Anoplophora glabripennis

The Asian longhorned beetle (ALB), Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae), is recognised as potentially one of the most damaging invasive insects in Europe and North America. International trade has increased the risk of accidental introduction of ALB. An eradication programme was initiated in Northeast Italy in June 2009, when an ALB infestation was discovered. The infestation was monitored by annual surveys of all host‐tree species growing in the eradication area. Infested trees were cut down and chipped. This study analyses the spatiotemporal distribution of infested trees for a 5‐year period from 2008 to 2012 using a generalised linear model approach. The results show that spread and infestation risk were significantly affected by (1) distance of suitable hosts from the nearest infested trees, (2) number of infested trees in the surroundings, and (3) annual variation. The significant differences in beetle dispersal between years reflect to some extent the onset of the eradication programme. The model allowed the estimation of arbitrary probability‐based management boundaries surrounding ALB‐infested trees. For example, the model estimated a 0.1% probability of attack on a suitable host tree 1 910 m from an existing attack.     

Spatio–temporal Variations in Post‐dispersal Seed Fate1

The post‐dispersal fate of Chrysophyllum lucentifolium (a canopy tree; Sapotaceae) seeds was analyzed in French Guiana over three consecutive years. Experiments using 750 thread‐marked seeds were performed to investigate seed removal, predation, and caching by terrestrial vertebrates on howler monkey (Alouatta seniculus) defecation sites, where clumps of intact C. lucentifolium seeds were observed. Year‐to‐year variations in seed fate during the peak fruiting period were considered in relation to overall fruit and seed resource availability estimated by a raked‐trail survey. The effect of two forest areas, which differed in soil and floristic composition, was examined with conspecific fruiting tree density as a covariant. Exclosure versus open treatment was used to discriminate small rodents (not larger than a spiny rat) from other vertebrates. The presence of fresh howler dung did not affect seed fate after 20 days as shown by comparisons between defecation sites and control during the first year. There was a significant effect of year on the percentage of seeds remaining after 20 days. Low seed removal in 1995 and 1996 (compared to 1997) corresponded to higher overall fruiting and higher fruiting of C. lucentifolium, or the presence of alternative resources for rodents. An effect of forest area was observed on the seed removal rate, which varied with years and protection. Comparatively, an effect of forest area on the percentage of seeds lost was observed in 1996 and an effect of treatment on the percentage of seeds eaten was seen in 1995. The mode of seed caching suggested that spiny rats were the main seed remover. Results of this study suggest that greater seedling recruitment may occur when large fruit crop and high howler dispersal co‐occur with a lower impact of rodents (i.e., when rodents are saturated by abundant and diversified fruit resources such as in 1995). Such event synchrony, however, is highly unpredictable after only three years of study.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.