Importance of bird traits for seed dispersal patterns of co‐fruiting trees in a patchy forest

Habitat fragmentation is globally one of the most important drivers of change in biodiversity. Seed dispersal by birds is crucial for tree regeneration in remnant patchy forests, yet how bird traits affect seed dispersal pattern is still poorly understood. We studied the extent to which bird traits affect seed‐removal networks and whether these traits affect seed deposition and seedling recruitment for 3 co‐fruiting tree species (Taxus chinensis, Cinnamomum bodinieri and Machilus thunbergii) in a patchy forest. A total of 17, 18 and 10 bird species were recorded foraging for seeds of T. chinensis, M. thunbergii and C. bodinieri, respectively. Frequency of bird visitation increased with tail length, wing length and body length. Furthermore, bird body length, bill length, body weight and wing length were important in the strength of the seed removal network. During foraging, 6 bird species exhibited different patterns of microhabitat utilization and their perching frequency increased with bird weight and tarsus length. As a consequence, frequency of habitat use, bird length and tarsus length were important in determining the number of deposited seeds. For seedling recruitment, seedling number increased with bird tarsus length and weight, but decreased with wing length. Overall, our results showed that various bird traits not only affected seed removal, but also influenced the subsequent processes of seed deposition and seedling distribution in a patchy forest. These results thus highlight the importance of large‐bodied birds for plant recruitment and point out the need to prioritize the protection and conservation of these birds in remnant patchy forests.     

Effect of frugivorous birds on the establishment of a naturally regenerating population of Chinese yew in ex situ conservation

The Chinese yew (Taxus chinensis) is catalogued as an endangered species in China because of the small size and senescent status of most populations. Its lack of natural regeneration is the most important reason for its endangered status. We investigated the regeneration of an ex‐situ conservation population, which was introduced into the Nanjing Botanical Garden Memorial Sun Yat‐Sen in the 1950s, and evaluated the role of frugivorous birds on the establishment of this regenerating population. Two hundred and thirteen individual yew seedlings and saplings were found on the hillside in 2006, and the nearest seedling was found more than 10 m away from mother trees. The spatial pattern of all seedlings and saplings occurred as a clumped distribution, which is typical for vertebrate‐dispersed plants. Six bird species were seen ingesting whole “fruits” at yew trees in the present study and 745 visits by avian frugivores were recorded. Red‐billed blue magpie (Urocissa erythrorhyncha), Chinese bulbul (Pycnonotus sinensis) and azure‐winged magpie (Cyanopica cyana) were the most frequent visitors over the two years of the study. Comparing the flights of departure and perching habitats of the three main bird species, we inferred that U. erythrorhyncha would be the most important disperser. This regeneration population has had good development over the past 20 or more years; frugivorous birds have removed seeds to the hillside every year until now, and seed germination and seedling growth continue to develop well under natural conditions. We suggest that the conservation system of the Chinese yew should be composed of yews, avian dispersers and habitats for seed germination and seedling growth.     

Endozoochory of frugivorous birds as potential dispersal of seed parasites from Pistacia chinesis: experimental evidence

The plant–disperser–fruit pest triads involve 3 interacting animals or groups (plants, vertebrates and seed parasites), and the dispersal of both seeds and seed parasites, which can both benefit from endozoochory via defecation or regurgitation by frugivorous vertebrates. However, we have very limited knowledge about the ecological and evolutionary consequences of these plant–disperser–fruit pest triads. Across central Northern China, several seed wasps (mainly Eurytoma plotnikov attack Pistacia chinensis fruits, and seed wasp larvae can develop, diapause and finally emerge as adults inside a seed during the following 1–3 years. In this study, we experimentally investigated whether frugivorous birds discriminated P. chinesis fruits with or without seed parasites, and whether bird endozoochory (by defecation or regurgitation) affected larval survival of seed parasites. The infestation rate by seed parasites was 37% of the P. chinesis fruit crop but with up to 48% of aborted fruits. We found that all 5 bird species can discriminate and then reject all unhealthy fruits (including aborted and insect‐infested). However, 4 of the 5 bird species, in particular bulbul species, consumed 15–41% of aborted and insect‐infested fruits as complementary food only when these unhealthy fruits were provided. Moreover, all larva of seed parasites remained alive after bird defecation or regurgitation. In conclusion, our study demonstrates that endozoochory by frugivorous birds could lead to potential dispersal of seed parasites of P. chinensis but with a very low probability.     

Effects of masting on seedling establishment of a rodent-dispersed tree species in a warm-temperate region,northern China

Masting is an evolutionary strategy used by plants to promote seed survival and/or seed dispersal under animal predation, but its effects on seedling establishment in field condition are rarely tested by long-term experiments incorporating combined effects of seed and animal abundance. Here, we tracked seed production, rodent-mediated seed dispersal, and seedling establishment in Armeniaca sibirica from 2005 to 2014 in a warm-temperate forest in northern China, and examined the effects of seed abundance and per capita seed availability on seed fate and seedling recruitment rate. Our results showed that seed abundance or per capita seed availability generally benefited the seedling recruitment of A. sibirica through increasing dispersal intensity, supporting predator dispersal hypothesis. However, seedling recruitment showed satiated or even dome-shaped association with per capita seed availability, suggesting the benefit to trees would be decreased when seed abundance were too high as compared to rodent abundance (a satiated effect). Our results suggest that the predator dispersal and satiation effects of masting on seedling recruitment can operate together in one system and conditionally change with seed and animal abundance.     

How seed defense and seed abundance predict dispersal and survival patterns in Camellia

Little is known about how seed defense and seed abundance interact with behavioral responses of seed dispersers to predict dispersal and survival dynamics in animal‐dispersed plants. By tracking the fate of individual seeds in Camellia stands with high and low seed abundance in Southwest China in 2007, we investigated the dispersal and survival of 2 high‐saponin Camellia species (Camellia oleifera and Camellia sinensis and 1 non‐saponin species (peanut Arachis hypogaea) as a control. Saponins in Camellia seeds are chemical compounds that act as seed defense. Our results were most consistent with the predictions based on the predator satiation hypothesis and the plant defense hypothesis. At the abundant Camellia stand (predators and dispersers were satiated), more Camellia seeds survived at the source but fewer were hoarded and survived at cache sites. At the sparse Camellia stand (predators and dispersers were not satiated), no Camellia seeds survived at the source, but more Camellia seeds were hoarded and survived at cache sites. Unlike Camellia seeds, no peanuts survived at the source at both stands, while more peanuts were hoarded and then survived at cache sites in the abundant Camellia stand compared to none at the sparse Camellia stand. In addition, the 2 Camellia species showed similar trends for seed fates across different dispersal stages. Our study indicates that the combined effects of seed abundance and seed defense, compared to their separate effects, provide a more accurate prediction for dispersal and survival patterns in animal‐dispersed Camellia species.     

Fluctuation in seed abundance has contrasting effects on the fate of seeds from two rapidly germinating tree species in an Asian tropical forest

The seed predator satiation hypothesis states that high seed abundance can satiate seed predators or seed dispersers, thus promoting seed survival. However, for rapidly germinating seeds in tropical forests, high seed abundance may limit dispersal as the seeds usually remain under parent trees for long periods, which may lead to high mortality due to rodent predation or fungal infestations. By tracking 2 species of rapidly germinating seeds (Pittosporopsis kerrii, family Icacinaceae; Camellia kissi, family Theaceae), which depend on dispersal by scatter‐hoarding rodents, we investigated the effects of seed abundance at the community level on predation and seed dispersal in the tropical forest of Xishuangbanna Prefecture, Southwest China. We found that high seed abundance at the community level was associated with delayed and reduced seed removal, decreased dispersal distance and increased pre‐dispersal seed survival for both plant species. High seed abundance was also associated with reduced seed caching of C. kissi, but it showed little effect on seed caching of P. kerrii. However, post‐dispersal seed survival for the 2 plant species followed the reverse pattern. High seed abundance in the community was associated with higher post‐dispersal survival of P. kerrii seeds, but with lower post‐dispersal survival of C. kissi seeds. Our results suggest that different plant species derive benefit from fluctuations in seed production in different ways.     

Does multiple seed loading in Blue Jays result in selective dispersal of smaller acorns?

Studies from both tropical and temperate systems show that scatter‐hoarding rodents selectively disperse larger seeds farther from their source than smaller seeds, potentially increasing seedling establishment in larger‐seeded plants. Size‐biased dispersal is evident in many oaks (Quercus) and is true both across and within species. Here, we predict that intraspecifc variation in seed size also influences acorn dispersal by the Blue Jay (Cyanocitta cristata Linnaeus), but in an opposite manner. Blue Jays are gape‐limited and selectively disperse smaller acorn species (e.g. pin oaks [Quercus palustris Münchh]), but often carry several acorns in their crop during a single dispersal event. We predict that jays foraging on smaller acorns will load more seeds per trip and disperse seeds to greater distances than when single acorns are carried in the bill. To test this, we presented free‐ranging Blue Jays with pin oak acorns of different sizes over a 2‐year period. In each of 16 experimental trials, we monitored the birds at a feeding station with remote cameras and determined the number of acorns removed and the distance acorns were dispersed when cached. Jays were significantly more likely to engage in multiple seed loading with smaller seeds in both years of the study. During the second year, these smaller acorns were dispersed farther than larger acorns, and during the first year, larger acorns were dispersed farther, revealing an inconsistent response to seed size during our study. We suggest that in some circumstances, multiple seed loading by Blue Jays may favor dispersal in some plant species.     

Effectiveness of seed dispersal by ants in a Neotropical tree

Seed dispersal and subsequent recruitment is the template on which forest regeneration takes place. Hence, considering the scale over which ecological processes occur is key for understanding the overall impact of various dispersal agents. To explore leafcutter ant (Atta colombica) dispersal effectiveness in space and time, seed movement and subsequent recruitment of a large‐seeded predominately vertebrate‐dispersed tree, Simarouba amara (Aubl. Simaroubaceae), was investigated on Barro Colorado Island, Panama. At each of 218 reproductive‐sized adults (≥20 cm diameter at breast height), presence or absence of a leafcutter ant colony was noted, with extensive checks for Atta activity taking place at or in close proximity to seed and seedling transects, which extended 4 cardinal directions for 30 m from each reproductive female tree (n= 74). Only at 2 S. amara trees were nests observed, and in these areas a dense S. amara seedling carpet was observed. Although nearby nest and dump sites might increase local S. amara recruitment in the short term, mortality at these sites is complete or nearly so. Hence, the seed dispersal effectiveness by leafcutter ants appears to be ephemeral and likely contributes inconsequentially to the long‐term recruitment and distribution patterns of the species. This finding highlights the importance of evaluating disperser effectiveness at ecologically relevant spatiotemporal scales.     

High rodent abundance increases seed removal but decreases scatter-hoarding and seedling recruitment along an elevational gradient

The distributions of small rodents in mountainous environments across different elevations can provide important information regarding the effects of climate change on the dispersal of plant species. However, few studies of oak forest ecosystems have compared the elevational patterns of sympatric rodent diversity, seed dispersal, seed bank, and seedling abundance. Thus, we tested the differences in the seed disperser composition and abundance, seed dispersal, seed bank abundance, and seedling recruitment for Quercus wutaishanica along 10 elevation levels in the Taihang Mountains, China. Our results provide strong evidence that complex asymmetric seed dispersal and seedling regeneration exist along an elevational gradient. The abundance of rodents had a significant negative correlation with the elevation and the seed removal rates peaked and then declined with increasing elevation. The seed removal rates were higher at middle and lower elevations than higher elevations but acorns were predated by 5 species of seed predators at middle and lower elevations, and thus, there was a lower likelihood of recruitment compared with those dropped beneath mother oaks at higher elevations. More importantly, the number of individual seeds in the seed bank and seedlings increased with the elevation, although dispersal services were reduced at sites lacking rodents. As conditional mutualists, the rodents could possibly act as antagonistic seed predators rather than mutualistic seed dispersers at low and middle elevations, thereby resulting in the asymmetric pattern of rodent and seedling abundance with increasing elevation to affect the community assembly and ecosystem functions on a large spatial scale.     

Exclusion of interspecific competition reduces scatter‐hoarding of Siberian chipmunk Tamias sibiricus: A field study

Although food availability and the abundance of seed predators have been postulated to affect seed dispersal, it is not clear how seed‐eating animals modify their scatter‐hoarding strategies in response to different levels of interspecific competition. We placed paired germinated and ungerminated acorns of Quercus mongolica on 30‐cm high platforms to exclude potential interspecific competition of the predominant larder hoarders Apodemus peninsulae and Myodes rufocanus, to investigate seed dispersal by a predominant scatter‐hoarder, Tamias sibiricus, in the field in north‐eastern China. Our results showed that T. sibiricus ate more acorns in situ in the absence of interspecific competition. In the presence of interspecific competition of A. peninsulae and C. rufocanus, however, more acorns were scatter‐hoarded by T. sibiricus. Regardless of interspecific competition, germination of acorns showed no significant effects on seed dispersal patterns, inconsistent with the “seed perishability hypothesis” that animals avoid hoarding seeds with high perishability. Exclusion of interspecific competition, though relatively increasing the per capita seed abundance, appears to reduce seed dispersal, scatter‐hoarding and seedling establishment. Therefore, we propose that moderate interspecific competition rather than competition exclusion may benefit seed scatter‐hoarding and seedling establishment.     

Seed traits and taxonomic relationships determine the occurrence of mutualisms versus seed predation in a tropical forest rodent and seed dispersal system

Although many studies have been carried out on plant–animal mutualistic assemblages, the roles of functional traits and taxonomy in determining both whether interactions involve mutualisms or predation and the structure of such assemblages are unclear. We used semi‐natural enclosures to quantitatively assess the interaction strengths between seeds of 8 sympatric tree species and 4 rodent species in a tropical forest in Xishuangbanna, Yunnan, Southwest China. We found 2 clusters of species in the seed–rodent network represented by 2 genera in the Fagaceae (Castanopsis, Lithocarpus). Compared to seeds of 3 Castanopsis species, seeds with heavy weight, hard coat or caloric content (including 3 Lithocarpus species) were eaten less and more frequently hoarded by rodents. In turn, hoarded seeds showed less predation and more mutualism with rodents. Our results suggest that seed traits significantly affected the hoarding behavior of rodents, and, consequently, the occurrence of mutualisms and predation as well as assemblage structure in the plant–animal seed dispersal system. Taxonomically‐related species with similar seed traits as functional groups belong to the same substructures in the assemblage. Our results indicate that both seed traits and taxonomic relationships may simplify thinking about seed dispersal systems by helping to elucidate whether interactions are likely to be dominated by predation or mutualism.     

Seed handling by primary frugivores differentially influence post‐dispersal seed removal of Chinese yew by ground‐dwelling animals

Seed handling by primary frugivores can influence secondary dispersal and/or predation of post‐dispersal seeds by attracting different guilds of ground‐dwelling animals. Many studies have focused on seeds embedded in feces of mammals or birds; however, less is known about how ground‐dwelling animals treat seeds regurgitated by birds (without pulp and not embedded in feces). To compare the effect of differential seed handling by primary dispersers on secondary seed removal of Chinese yew (Taxus chinensis var. mairei), we conducted a series of exclosure experiments to determine the relative impact of animals on the removal of defecated seeds (handled by masked palm civet), regurgitated seeds (handled by birds) and intact fruits. All types of yew seeds were consistently removed at a higher rate by rodents than by ants. Regurgitated seeds had the highest removal percentage and were only removed by rodents. These seeds were probably eaten in situ without being secondarily dispersed. Defecated seeds were removed by both rodents and ants; only ants might act as secondary dispersers of defecated seeds, whereas rodents ate most of them. We inferred that seeds regurgitated by birds were subjected to the highest rates of predation, whereas those dispersed in the feces of masked palm civets probably had a higher likelihood of secondary dispersal. Seeds from feces attracted ants, which were likely to transport seeds and potentially provided a means by which the seeds could escape predation by rodents. Our study highlighted that primary dispersal by birds might not always facilitate secondary dispersal and establishment of plant populations.     

Cultivated walnut trees showed earlier but not final advantage over its wild relatives in competing for seed dispersers

Little is known about seeding regeneration of cultivated trees compared to wild relatives in areas where seed dispersers are shared. Here, we investigated the differences in seed fates of cultivated walnut (Juglans regia) and wild Manchurian walnut (Juglans mandshurica) trees under rodent predation and dispersal. J. regia seeds have higher nutritional value (large size, mass and kernel mass) and lower mechanical defensiveness (thin endocarp) than J. mandshurica seeds. We tracked seeds of J. regia and J. mandshurica under both enclosure and field conditions to assess differences in competing for seed dispersers of the two co‐occurring tree species of the same genus. We found that rodents preferred to harvest, eat and scatter‐hoard seeds of J. regia as compared to those of J. mandshurica. Seeds of J. regia were removed and scatter‐hoarded faster than those of J. mandshurica. Caches of J. regia were more likely to be rediscovered by rodents than those of J. mandshurica. These results suggest that J. regia showed earlier dispersal fitness but not the ultimate dispersal fitness over J. mandshurica in seeding regeneration under rodent mediation, implying that J. regia has little effect on seeding regeneration of J. mandshurica in the field. The effects of seed traits on seed dispersal fitness may vary at different dispersal stages under animal mediation.     

Scatter‐hoarding rodents as secondary seed dispersers of a frugivore‐dispersed tree Scleropyrum wallichianum in a defaunated Xishuangbanna tropical forest,China

Local extinction or population decline of large frugivorous vertebrates as primary seed dispersers, caused by human disturbance and habitat change, might lead to dispersal limitation of many large‐seeded fruit trees. However, it is not known whether or not scatter‐hoarding rodents as secondary seed dispersers can help maintain natural regeneration (e.g. seed dispersal) of these frugivore‐dispersed trees in the face of the functional reduction or loss of primary seed dispersers. In the present study, we investigated how scatter‐hoarding rodents affect the fate of tagged seeds of a large‐seeded fruit tree (Scleropyrum wallichianum Arnott, 1838, Santalaceae) from seed fall to seedling establishment in a heavily defaunated tropical forest in the Xishuangbanna region of Yunnan Province, in southwest China, in 2007 and 2008. Our results show that: (i) rodents removed nearly all S. wallichianum seeds in both years; (ii) a large proportion (2007, 75%; 2008, 67.5%) of the tagged seeds were cached individually in the surface soil or under leaf litters; (iii) dispersal distance of primary caches was further in 2007 (19.6 ± 14.6 m) than that in 2008 (14.1 ± 11.6 m), and distance increased as rodents recovered and moved seeds from primary caches into subsequent caching sites; and (iv) part of the cached seeds (2007, 3.2%; 2008, 2%) survived to the seedling stage each year. Our study suggests that by taking roles of both primary and secondary seed dispersers, scatter‐hoarding rodents can play a significant role in maintaining seedling establishment of S. wallichianum, and are able to at least partly compensate for the loss of large frugivorous vertebrates in seed dispersal.     

Endozoochorous seed dispersal by golden snub-nosed monkeys in a temperate forest

Seed dispersal is essential for plant recruitment and the maintenance of biodiversity. Colobine monkeys are primarily folivorous, but they also consume fruits and are often assumed to be seed predators. Although they are known to be epizoochorous seed dispersers, their role as endozoochorous seed dispersers needs reassessment. We examined potential endozoochory in golden snub-nosed monkeys (Rhinopithecus roxellana) at Dalongtan in Shennongjia National Park, central China, by assessing potential germination of ingested seeds (n = 1806, 9 species) from fecal samples. Intact seeds were in almost all fecal samples (ranging from 5–130 seeds), and ingested seeds were from small seeded species (seed width <4.5 mm). The 2 most abundant species were Actinidia arguta (73%) and Rosa caudata (15%). The fruits of A. arguta were unripe when ingested (i.e. effective seed predation) and the ingested seeds did not germinate in the trials. Therefore, ingestion of unripe seeds does not lead to effective seed dispersal. However, germination rates of defecated R. caudata (9%) were greater than control seeds (6% and 0%), demonstrating potential endozoochorous seed dispersal. Thus, colobine monkeys do indeed disperse mainly small-seeded from multi-seeded fruits through potential endozoochory and this process enhances the recruitment of seedlings.     

Trait‐mediated seed predation,dispersal and survival among frugivore‐dispersed plants in a fragmented subtropical forest,Southwest China

By tracking the fate of individual seeds from 6 frugivore‐dispersed plants with contrasting seed traits in a fragmented subtropical forest in Southwest China, we explored how rodent seed predation and hoarding were influenced by seed traits such as seed size, seed coat hardness and seed profitability. Post‐dispersal seed fates varied significantly among the 6 seed species and 3 patterns were witnessed: large‐seeded species with a hard seed coat (i.e. Choerospoadias axillaries and Diospyros kaki var. silvestris) had more seeds removed, cached and then surviving at caches, and they also had fewer seeds predated but a higher proportion of seeds surviving at the source; medium‐sized species with higher profitability and thinner seed coat (i.e. Phoebe zhennan and Padus braohypoda) were first harvested and had the lowest probability of seeds surviving either at the source or at caches due to higher predation before or after removal; and small‐seeded species with lower profitability (i.e. Elaeocarpus japonicas and Cornus controversa) had the highest probability of seeds surviving at the source but the lowest probability of seeds surviving at caches due to lower predation at the source and lower hoarding at caches. Our study indicates that patterns of seed predation, dispersal and survival among frugivore‐dispersed plants are highly determined by seed traits such as seed size, seed defense and seed profitability due to selective predation and hoarding by seed‐eating rodents. Therefore, trait‐mediated seed predation, dispersal and survival via seed‐eating rodents can largely affect population and community dynamics of frugivore‐dispersed plants in fragmented forests.     

Digestive efficiency of indigenous and invasive avian species fed fruit of invasive alien plants in South Africa

Many highly invasive plant species produce fleshy fruit that are consumed and dispersed by frugivorous birds. However, little is known about assimilation efficiency of invasive fruit by indigenous and invasive avian species. We investigated whether indigenous Knysna (Tauraco corythaix) and Purple-crested (Gallirex porphyreolophus) Turacos and invasive alien Rose-ringed Parakeets (Psittacula krameri) met their energy demands when fed fruits of four fleshy- fruited invasive alien plant species: Solanum mauritianum, Cinnamomum camphora, Psidium guajava and Morus alba. Birds were fed single-fruit diets for two consecutive days and energetic parameters were calculated for all fruit diets. Our results showed that generally both invasive and indigenous avian species managed to gain their daily energy requirements from fruits of the four respective invasive plants, suggesting that they can meet their energetic demands by feeding on them only. The exception was P. krameri, which did not feed on S. mauritianum fruit. These findings may explain why fruits of invasive alien plants are attractive to most avian frugivores and highlight the role of avian frugivores in their dispersal.     

The function of the hygroscopic awn of Themeda triandra 1

The hygroscopic awn of species such as Themeda triandra is widely believed to be a device ensuring the early burial of its seed. This belief has been extensively tested and no tendency for such self‐burial has been found. It has been demonstrated that the propelling action of the awn does cause the seed to move along the surface and its possible function as a dispersal mechanism is tentatively suggested.     

有关白颈长尾雉雄性偏倚扩散的遗传学证据

一种性别的个体倾向于比另一性别的个体扩散到离出生地较远的地方,这种现象称为性别偏倚扩散.性别偏倚扩散现象在脊椎动物中很常见,对种群结构和种群动态有着非常重要的作用.许多研究已经证明了一夫一妻制鸟类的扩散模式,然而有关一夫多妻制鸟类的研究和数据还是很少.白颈长尾雉(Syrmaticus ellioti)是中国特有的濒危物种.本研究中,我们使用多态性DNA微卫星位点(105个个体和7个位点)和线粒体DNA控制区序列(63个个体)来研究白颈长尾雉的性别偏倚扩散模式.传统的观点认为,鸟类中雌性个体比雄性个体更具扩散性.与此不同的是,本文中两种遗传标记所得的遗传信息都表明,白颈长尾雉的扩散模式是雄性偏倚的.我们认为,在鸡形目中,不具群集展示行为的一夫多妻制物种更加倾向于雄性偏倚扩散模式.该推论与基于白颈长尾雉一夫多妻的婚配体制所得到的结果是相一致的.     

Germination of One-Seed Juniper Seeds Distributed by Different Frugivore Groups

One-seed juniper (Juniperus monosperma [Englem.] Sarg.) encroachment into grassland habitats is facilitated by a diverse group of frugivores. To test seed germination after gut passage, we collected pellets or scats containing mature seeds from four frugivore groups: passerine (perching) birds, lagomorphs (hares and rabbits), mesocarnivores (coyotes and foxes), and porcupines. For comparison of germination success, we also evaluated seeds from intact, nondigested one-seed juniper cones. We conducted germination trials under three different scenarios: 1) imbibed bare seeds (a measure of potential germination); 2) seeds still encased within a frugivore deposited pellet or scat matrix or still in a nondigested cone; and 3) matrix-free seeds that had gone through the matrix germination trials and then were freed from the matrix (a simulation of seeds that become dislodged from scats or pellets). In the bare seed trials, germination was highest (70.8% ± 7.4%) for seeds digested by mesocarnivores and was 63.9% ± 5.7% for porcupines, which was higher (P < 0.05) than germination of seeds from bird pellets (33.3% ± 7.7%). Germination of seeds from all frugivores, including lagomorphs (51.4% ± 7.2%), was higher (P < 0.05) than for seeds encased in cones (9.7% ± 3.2%). The germination percentage of seeds still encased in frugivore pellets or scats was highest for birds (40.1 ± 4.2) and was significantly lower (P < 0.05) for other frugivores. Seeds freed from all frugivore deposition matrices showed a second pulse of germination activity, especially seeds from lagomorph pellets. Germination success differences can influence the nature of one-seed juniper woodland infill and encroachment into bordering grasslands, which can lead to loss of soil health, lower grass biomass production, and a diminishment of habitat quality for grassland-dependent wildlife species.