Water and fish select for fleshy fruits in tropical wetland forests

Adjacent floodplain and upland tropical forests experience the same temperature and precipitation regimes, but differ substantially in plant species composition and biotic interactions because of extensive flooding. We hypothesize that flooded forests filter fruiting traits linked to seed dispersal by water and fishes, such that selection by water and fish led to (1) trees that synchronize the timing of fruiting with annual floods, and (2) the evolution of fleshy tissues on fruits to improve buoyancy and increase fruit consumption rates by fish. To test this hypothesis, we compared plant communities in seasonally flooded forests and adjacent upland forest in terms of fruiting phenology, the frequency of trees bearing fleshy fruit, and the role of fleshy tissues in buoyancy and seed viability. Beta‐diversity in this system is high, with significant differences in species composition across habitats. As predicted, the production of ripe fleshy fruits was significantly greater in flooded than upland forests during the flood season. Furthermore, we found that trees with fleshy fruit were significantly more abundant in flooded forests even though species richness of fleshy fruit‐bearing trees was proportionally similar in flooded and upland forests. Additionally, fleshy pulp increased buoyancy. Likewise, time afloat decreased for denser fruit and those with high seed to pulp ratios. In concert, these results suggest that fleshy fruits in Neotropical floodplain forests facilitated hydrochory and ichthyochory. Once established, water and fish became important agents of selection on fruiting traits.     

Frugivory and Seed Dispersal by Northern Pigtailed Macaques (Macaca leonina), in Thailand

Tropical rain forest conservation requires a good understanding of plant–animal interactions. Seed dispersal provides a means for plant seeds to escape competition and density-dependent seed predators and pathogens and to colonize new habitats. This makes the role and effectiveness of frugivorous species in the seed dispersal process an important topic. Northern pigtailed macaques (Macaca leonina) may be effective seed dispersers because they have a diverse diet and process seeds in several ways (swallowing, spitting out, or dropping them). To investigate the seed dispersal effectiveness of a habituated group of pigtailed macaques in Khao Yai National Park, Thailand, we examined seed dispersal quantity (number of fruit species eaten, proportion in the diet, number of feces containing seeds, and number of seeds processed) and quality (processing methods used, seed viability and germination success, habitat type and distance from parent tree for the deposited seeds, and dispersal patterns) via focal and scan sampling, seed collection, and germination tests. We found thousands of seeds per feces, including seeds up to 58 mm in length and from 88 fruit species. Importantly, the macaques dispersed seeds from primary to secondary forests, via swallowing, spitting, and dropping. Of 21 species, the effect of swallowing and spitting was positive for two species (i.e., processed seeds had a higher % germination and % viability than control seeds), neutral for 13 species (no difference in % germination or viability), and negative (processed seeds had lower % germination and viability) for five species. For the final species, the effect was neutral for spat-out seeds but negative for swallowed seeds. We conclude that macaques are effective seed dispersers in both quantitative and qualitative terms and that they are of potential importance for tropical rain forest regeneration.     

Gut passage effect of the introduced red‐whiskered bulbul (Pycnonotus jocosus) on germination of invasive plant species in Mauritius

In Mauritius, many of the worst invasive plant species have fleshy fruits and rely on animals for dispersal. The introduced red‐whiskered bulbul (Pycnonotus jocosus) feeds on many fleshy‐fruited species, and often moves from invaded and degraded habitats into higher quality native forests, thus potentially acting as a mediator of continued plant invasion into these areas. Furthermore, gut passage may influence seed germination. To investigate this, we fed fleshy fruits of two invasive plant species, Ligustrum robustum and Clidemia hirta, to red‐whiskered bulbuls. Gut passage times of seeds were recorded. Gut‐passed seeds were sown and their germination rate and germination success compared with that of hand‐cleaned seeds, as well as that of seeds in whole fruits. Gut passage and hand‐cleaning had significant positive effects on germination of both species. Gut‐passed seeds of both C. hirta and L. robustum germinated faster than hand‐cleaned seeds. However, for L. robustum, this was only true when compared with hand‐cleaned seeds with intact endocarp; when compared with hand‐cleaned seeds without endocarp, there was no difference. For overall germination success, there was a positive effect of gut passage for C. hirta, but not for L. robustum. For both C. hirta and L. robustum, no seeds in intact fruits geminated, suggesting that removal of pulp is essential for germination. Our results suggest that, first, the initial invasion of native forests in Mauritius may not have happened so rapidly without efficient avian seed dispersers like the red‐whiskered bulbul. Second, the bulbul is likely to be a major factor in the continued re‐invasion of C. hirta and L. robustum into weeded and restored conservation management areas.     

Germination of Geonoma brevispatha (Arecaceae) in laboratory and its relation to the palm spatial distribution in a swamp forest

Geonoma brevispatha is a clonal palm species commonly found as tussocks in swamp and riparian forests in SE Brazil, occurring in the very transition between flooded pits/channels and dry mounds. Aiming to verify whether the germination responses could explain the spatial pattern of the palm tussocks in swamp forests, we investigated the germination in relation to the main environmental factors to which the seeds are subjected in their natural habitat. The seeds are recalcitrant since germination percentage is reduced with loss of moisture content. Germination did not differ on moist filter paper, semi-immersion (water up to half of the seed diameter) or complete immersion (water up to double of the seed diameter). Seeds did not germinate when stored in a closed bottle with water (that is, under low oxygen concentration), but remained viable for up to 8 weeks under this condition. The capacity to germinate in water with minimum oxygen availability may be considered ecologically advantageous in a swamp habitat. On the other hand, the capacity to germinate in moist filter paper indicates that, in natural conditions, the seed can germinate, provided there is a critical minimum of water availability. We concluded that the germination responses per se are not enough to explain the palm tussocks spatial pattern, but they are very well tuned with the conditions of the transitional zone between channels/pits (ever flooded micro-sites) and mounds (well-drained micro-sites), in which the species occurs in the swamp forest.     

Seed Removal Rates Under Isolated Trees and Continuous Vegetation in Semiarid Thornscrub

Clearing native vegetation to increase the amount of land available for agriculture in northeastern Mexico has left remnants ranging in size from fragments of continuous vegetation to isolated individual trees. These provide valuable opportunities for restoring larger areas of native vegetation. We explored whether fragmentation of Tamaulipan thornscrub affects the removal of seeds from 12 woody species that encompass a range of sizes and dispersal mechanisms. We tested whether (1) seed removal rates under isolated trees were higher than under continuous vegetation; (2) dispersal structures, such as fleshy pericarps, made some seeds more attractive to seed removers; and (3) microenvironmental variation affected seed removal rates. Seeds were placed under canopies of Texas ebony (Ebenopsis ebano) and Mesquite trees (Prosopis laevigata). Seed removal trials were conducted three times, each trial lasting 30 days. Most seeds were removed in all trials by the end of one month. Seed removal rate was slower under isolated trees. In general, fleshy fruits were removed faster than other fruits; whole fruits and fleshy tissue were removed faster than depulped seeds. In species with fleshy pericarps, acid washing of seeds, to simulate seeds processed in the digestive tract of dispersers, reduced the seed removal rates, suggesting that it would be a good pre‐treatment for restoration efforts.     

A water drop-shaped slingshot in plants: geometry and mechanics in the explosive seed dispersal of Orixa japonica (Rutaceae)

Background and AimsIn angiosperms, many species disperse their seeds autonomously by rapid movement of the pericarp. The fruits of these species often have long rod- or long plate-shaped pericarps, which are suitable for ejecting seeds during fruit dehiscence by bending or coiling. However, here we show that fruit with a completely different shape can also rely on pericarp movement to disperse seeds explosively, as in Orixa japonica.MethodsFruit morphology was observed by hard tissue sectioning, scanning electron microscopy and micro-computed tomography, and the seed dispersal process was analysed using a high-speed camera. Comparisons were made of the geometric characteristics of pericarps before and after fruit dehiscence, and the mechanical process of pericarp movement was simulated with the aid of the finite element model.Key ResultsDuring fruit dehydration, the water drop-shaped endocarp of O. japonica with sandwich structure produced two-way bending deformation and cracking, and its width increased more than three-fold before opening. Meanwhile the same shaped exocarp with uniform structure could only produce small passive deformation under relatively large external forces. The endocarp forced the exocarp to open by hygroscopic movement before seed launching, and the exocarp provided the acceleration for seed launching through a reaction force.ConclusionsTwo layers of water drop-shaped pericarp in O. japonica form a structure similar to a slingshot, which launches the seed at high speed during fruit dehiscence. The results suggest that plants with explosive seed dispersal appear to have a wide variety of fruit morphology, and through a combination of different external shapes and internal structures, they are able to move rapidly using many sophisticated mechanisms.     

Seed dispersal by the Florida box turtle (<Emphasis Type="Italic">Terrapene carolina bauri</Emphasis>) in pine rockland forests of the lower Florida Keys,United States

Seed dispersal by animals is one of the most important plant-animal mutualisms, but saurochory, the dispersal of seeds by reptiles, has received little attention. We investigated the role of the Florida box turtle (Terrapene carolina bauri) as a seed dispersal agent in pine rockland forests of the lower Florida Keys and examined the effect of turtle digestion on seed germination. We obtained seeds of 11 species with fleshy fruits and 2 species with non-fleshy fruits (a grass and legume) from the feces of 145 box turtles collected on Key Deer National Wildlife Refuge from 1999 to 2000. We planted the seeds of nine species and germination percentage (percentage of seeds that germinated during the experiment) varied from 10% to 80%. Comparative germination experiments were conducted with Thrinax morrissii, Serenoa repens, and Byrsonima lucida. We compared the germination percentage and germination rate (number of days from planting to seedling emergence) of seeds from three treatments (seeds recovered from feces, control seeds with pulp, and control seeds without pulp) and continued these experiments for up to 2 years. Passage through the box turtle digestive tract greatly enhanced the germination percentage and germination rate of S. repens, but decreased the germination percentage of B. lucida and T. morrissii, and decreased germination rate for T. morrissii. Subsequent destructive seed viability tests revealed that many ungerminated T. morrissii seeds remained viable, suggesting long-term seed dormancy may occur, even after passage through the turtle digestive system. In addition, the proportion of ungerminated seeds which remained viable was greater for seeds recovered from turtle feces than from control seeds with pulp. Furthermore, removal of fleshy pulp either manually or by the turtle digestive system may allow T. morrissii to escape insect predation.     

Germination in four low-várzea tree species of Central Amazonia

Trees of Central Amazonian white-water (várzea) forests are highly adapted to the annual inundations, which can last up to 7 months every year. Many trees synchronize fruit production to the period of highest water levels of the rivers, and hydrochory is especially common in species that colonize the low-lying flood-levels flooded for longer periods. The effect of the contact of diaspores with the river water is controversially discussed in literature. While many studies describe that flooding breaks the dormancy in seeds of many várzea tree species and is necessary for germination, other studies mention that seed buoyancy and/or submergence have negative effects on germination. Therefore, the present study was designed in order to test experimentally how seed buoyancy and seed submergence affect germination in four várzea tree species of the low-lying flood-levels. The tested species with buoyant seeds were Salix martiana and Pseudobombax munguba, those with submerged seeds Laetia corymbulosa and Vitex cymosa. 50 seeds from each species were (a) placed in water during a period of 15 days and afterwards moved to várzea substrate, thus simulating seed buoyancy and/or submergence in the natural environment, and (b) directly placed in várzea substrate, with four repetitions, respectively. Three species showed significantly higher percentages of germination in the flooded seeds than in the non-waterlogged seeds, while fruit-fibre involved seeds of P. munguba showed an opposite trend. In L. corymbulosa, germination initiated earlier in the submerged than in the control seeds, whereas there was no difference in the start of germination between waterlogged and non-waterlogged seeds of the other species. From buoyant seeds of P. munguba and S. martiana, seedlings with entirely formed cotyledons were developed while still in water. These seedlings were characterized by morphological differences in comparison to seedlings originating from non-waterlogged seeds and could not protrude the root into the soil (i.e. establish) when placed in the substrate. It is likely that the seed involving fruit-fibres contribute to long-distance dispersal in these species in the natural environment, and to stabilize seedlings when diaspores land on substrate. Concluding, contact with the river water did not disturb but on the contrary enhanced germination in the four studied species.     

Biogeography of seed-dispersal syndromes, life-forms and seed sizes among woody rain-forest plants in Australia's subtropics

Aim To enhance our understanding of the evolutionary interactions between seed‐dispersal syndromes, life‐forms, seed size, and habitat characteristics by studying their association with the regional‐scale distributions of subtropical rain‐forest plants in the context of climatic gradients. Location South‐east Queensland, subtropical eastern Australia (152° E, 26° S). Methods We classified 250 rain‐forest sites into six floristic site‐groups based on their woody plant composition. The resulting classification was strongly associated with variation in rainfall. The distribution of species across the floristic site‐groups was used to assign 568 species to seven habitat classes (one class for ‘widespread’ species, with all other species classified according to the site‐group within which they were most frequent). Species were also classified for three other categorical life‐history factors: three dispersal syndromes based on diaspore morphology (fleshy, wind‐assisted, and unadorned); four life‐forms (trees, shrubs and small trees, tall climbers, and short and shrubby climbers); and four seed‐diameter classes (< 3 mm, ≥ 3 and < 4.5 mm, ≥ 4.5 and < 7 mm, and ≥ 7 mm). We used a basic comparative approach augmented by simple phylogenetically constrained comparisons to assess association between dispersal syndrome, seed size, life‐form, and habitat class. Results Across the rain forests of south‐east Queensland, the proportion of species with fleshy diaspores or of large stature increases with rainfall. High‐rainfall sites also have larger average seed sizes, but the difference in average seed size between high‐ and low‐rainfall sites is small compared with variation within sites. Among species, those with fleshy fruit tend to have larger seeds and to favour high‐rainfall sites. Very few small trees produce diaspores adapted for wind‐assisted dispersal. On average, species with unadorned diaspores have smaller seeds than those with fleshy diaspores. However, within sites, species with unadorned and fleshy diaspores have similar average seed sizes, and some species with unadorned diaspores from high‐rainfall habitats have extremely large seeds. Main conclusions Commonly observed associations between fleshy fruit, larger plants, larger seeds, and productive habitats are apparent within the rain‐forest flora of south‐east Queensland. However, these associations are generally weak and involve complex interactions. For example, the strong tendency for species with fleshy fruit to have larger seeds than those with unadorned diaspores concealed a significant group of species from wetter forests that produce extremely large seeds and unadorned diaspores. The most widespread species in this study tend to be large plants (particularly robust lianes) and to produce fleshy fruit, but they tend not to have relatively large seeds. The association between large seeds, large plants, fleshy fruit and productive habitats is discussed as part of an evolutionary strategy favouring fitness in populations close to carrying capacity. We review some problems with focusing on establishment chances per seed as the driver towards association between large seeds, large plants and productive rain‐forest habitats (the difficult‐establishment hypothesis). Instead we suggest that production of large, short‐lived seeds by long‐lived plants in temporally stable, closed habitats may reflect the limited evolutionary potential for strategies enhancing colonization (e.g. producing large numbers of dormant seeds), thus allowing the establishment benefits of large seeds greater selective influence (the slow‐replacement hypothesis). The association of fleshy fruit with large seeds probably reflects the difficulty of dispersing large seeds by other means (the difficult‐dispersal hypothesis).     

A role for fruit structure in seed survival and germination of Swartzia langsdorffii Raddi beyond dispersal

• Diaspore structure has been hypothesised to play a role in seed viability and/or germination of recalcitrant seeds, especially for Swartzia langsdorffii. Thus, this work aims to (i) investigate the in situ contribution of pericarp and aril on seed viability and germination, and (ii) identify morphoanatomical traits of S. langsdorffii diaspores that allow its desiccation‐sensitive seeds to remain viable.
• The role of the pericarp and aril in seed survival and germination was investigated by placing the whole fruit, whole seeds (arillate seed) and bare seeds (without aril) in soil in the forest understorey, assessing germination, emergence, dead, firm and predated seeds, and water content of pericarps, arils and seeds. Correlation analysis was performed between environmental variables and physiological parameters. Histochemical features of diaspores were also investigated.
• Pericarp water content fell after several months, while the aril maintained its water content. Seeds did not lose water even without the presence of the pericarp and aril. However, presence of the pericarp promoted seed water content, viability and germination long after dispersal. The embryo had a thickened outer periclinal cell wall.
• Pericarp and aril are not essential to prevent water loss in seeds, but do help to retain seed moisture, favouring viability maintenance and promoting germination during the rainy season. Morphoanatomical features of seeds are suggested as main factors that reduce water loss. Survival of these desiccation‐sensitive seeds upon dispersal during the dry season appears to be facilitated by multiple diaspore features that prevent viability loss.

     

High-quality seed dispersal by fruit-eating fishes in Amazonian floodplain habitats

Seed dispersal is a critical stage in the life history of plants. It determines the initial pattern of juvenile distribution, and can influence community dynamics and the evolutionary trajectories of individual species. Vertebrate frugivores are the primary vector of seed dispersal in tropical forests; however, most studies of seed dispersal focus on birds, bats and monkeys. Nevertheless, South America harbors at least 200 species of frugivorous fishes, which move into temporarily flooded habitats during lengthy flood seasons and consume fruits that fall into the water; and yet, we know remarkably little about the quality of seed dispersal they effect. We investigated the seed dispersal activities of two species of large-bodied, commercially important fishes (Colossoma macropomum and Piaractus brachypomus, Characidae) over 3 years in Pacaya-Samiria National Reserve (Peru). We assessed the diet of these fishes during the flood season, conducted germination trials with seeds collected from digestive tracts, and quantified fruit availability. In the laboratory, we fed fruits to captive Colossoma, quantified the proportion of seeds defecated by adult and juvenile fish, and used these seeds in additional germination experiments. Our results indicate that Colossoma and Piaractus disperse large quantities of seeds from up to 35% of the trees and lianas that fruit during the flood season. Additionally, these seeds can germinate after floodwaters recede. Overexploitation has reduced the abundance of our focal fish species, as well as changed the age structure of populations. Moreover, older fish are more effective seed dispersers than smaller, juvenile fish. Overfishing, therefore, likely selects for the poorest seed dispersers, thus disrupting an ancient interaction between seeds and their dispersal agents. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Role of trichomes and pericarp in the seed biology of the desert annual Lachnoloma lehmannii (Brassicaceae)

In many angiosperms, the fruit rather than the seed is the dispersal/germination unit, and this is the case with Lachnoloma lehmannii, a desert annual ephemeral in central southwestern Asia with indehiscent nonmucilaginuous silicles covered with trichomes. The primary aim of this study was to assess the role of trichomes and pericarp in dispersal, anchorage of diaspores, and seed germination of this species. Mature silicles are dispersed by wind and gravity, and trichomes not only significantly increased their dispersal distance, adherence to sandy soil particles, mass of water imbibed and moisture content, but also decreased the rate of water loss and moisture content of seeds. A significantly higher percentage of seeds within silicles than of isolated seeds retained viability after exposure to 60 °C for 24 h. Seed dormancy is due to the pericarp and to nondeep physiological dormancy, as shown by the increase in germination percentage of isolated seeds following dry storage and treatment with GA₃. Removal of pericarp increased germination of 6-month-old seeds from 0 to 80–90 %, and leachate from both pericarp and trichomes significantly inhibited germination of isolated seeds. Ninety-five percent of seeds within silicles buried in soil for 2 years were viable, but only 28 % of them germinated in light at 15/2 °C; thus L. lehmannii forms a persistent soil seed bank. The pericarp and its trichomes may maximize plant fitness by determining the settlement location of silicles, thus helping to ensure that seeds germinate during the cool season for seedling survival in the desert environment.     

Processes regulating the invasion of European buckthorn (Rhamnus cathartica) in three habitats of the northeastern United States

Invasion by exotic plants often is restricted by processes, such as seed predation, acting on early life-history stages; however, the relative importance of these processes might vary among habitats. Modern human land use has created a mosaic of habitats in many landscapes, including the landscape of the northeastern United States. European buckthorn (Rhamnus cathartica) is an exotic plant that has achieved varying success in North American habitats. We studied dispersal, seed survival, germination, and seedling survival of buckthorn populations at four plots in each of sugar maple (Acer saccharum) forests, old fields, and abandoned conifer plantations in central New York State. Dispersal was low in maple forests, as evidenced by low collection rates of R. cathartica seeds in seed traps. Rates of post-dispersal seed predation were highest in maple forests and lowest in old fields, suggesting greater use of maple forests by granivorous rodents. Germination rates did not vary among seeds planted in soils of these habitats when studied in the laboratory despite differences in soil pH. Survival of transplanted seedlings was low in maple forests relative to old fields and plantations. Buckthorn invasion of old fields and abandoned plantations was not strongly constrained by factors we considered, and the buckthorn populations in these habitats were large. A combination of low dispersal by frugivores, low seed survival due to predation, and low seedling survival due to dim light conditions apparently prevents R. cathartica from invasion of intact maple forests of our area. Native fauna and canopy closure may act synergistically to reduce success of invasive plants in natural habitats.     

Seed flotation and germination of salt marsh plants: The effects of stratification, salinity, and/or inundation regime

We examined the effects of cold stratification and salinity on seed flotation of eight salt marsh species. Four of the eight species were tested for germination success under different stratification, salinity, and flooding conditions. Species were separated into two groups, four species received wet stratification and four dry stratification and fresh seeds of all species were tested for flotation and germination. Fresh seeds of seven out of eight species had flotation times independent of salinity, six of which had average flotation times of at least 50 d. Seeds of Spartina alterniflora and Spartina patens had the shortest flotation times, averaging 24 and 26 d, respectively. Following wet stratification, the flotation time of S. alterniflora seeds in higher salinity water (15 and 36 ppt) was reduced by over 75% and germination declined by more than 90%. Wet stratification reduced the flotation time of Distichlis spicata seeds in fresh water but increased seed germination from 2 to 16% in a fluctuating inundation regime. Fresh seeds of Iva frutescens and S. alternflora were capable of germination and therefore are non-dormant during dispersal. Fresh seeds of I. frutescens had similar germination to dry stratified seeds ranging 25-30%. Salinity reduced seed germination for all species except for S. alterniflora. A fluctuating inundation regime was important for seed germination of the low marsh species and for germination following cold stratification. The conditions that resulted in seeds sinking faster were similar to the conditions that resulted in higher germination for two of four species.     

Peduncles elicit large-mammal endozoochory in a dry-fruited plant

Background and Aims

Plants have evolved a variety of seed dispersal mechanisms to overcome lack of mobility. Many species embed seeds in fleshy pulp to elicit endozoochory, i.e. disseminating seed through the animal gut. In contrast to well-studied fleshy fruited plants, dry-fruited plants may exploit this dispersal mutualism by producing fleshy appendages as a nutritional reward to entice animals to swallow their diaspores, but this has been little studied. In this study, it is hypothesized that these accessory fruits represent co-adaptations facilitating the syndrome of mammalian endozoochorous dispersal.

Methods

Field observations (focal tree watches, faecal surveys and fruiting phenology) with experimental manipulations (examination of seed germination and feeding trials) were conducted over 2 years in a native population of the raisin tree, Hovenia dulcis, which produces enlarged, twisted brown peduncles with external black seeds, in central China.

Key Results

Birds were not observed to swallow seeds or carry infructescences away during 190 h of focal tree watches. However, H. dulcis seeds were detected in 247 faecal samples, representative of two herbivore and four carnivore mammalian species. Feeding trials revealed that peduncles attracted mammals to consume the entire infructescence, thereby facilitating effective seed dispersal. The germination rate of egested seeds proved higher than that of unconsumed seeds. It was also noted that this mutualism was most vulnerable in degraded forest.

Conclusions Hovenia dulcis

peduncle sets are confirmed to adapt primarily to mammalian endozoochory, a mutualistic association similar in function to fleshy pulp or foliage. This demonstrates that plant organ systems can be adapted to unique mutualisms that utilize animal dispersal agents. Such an ecological role has until now been attributed only to bird epizoochory. Future studies should consider more widely the putative role of peduncle sets and mammalian endozoochory as a dispersal mechanism, particularly for those plants that possess relatively large accessory fruits.     

Seed dimorphism and germination behaviour in Salicornia patula

One of the annual species of the genus Salicornia from the Mediterranean coast, Salicornia patula presents a seed dimorphism. This morphological seed dimorphism is linked with a physiological ‘dimorphism’ with regard to some requirements for germination. Central seeds, in the inflorescence, are larger, are dispersed attached to the perianth, have no light requirement for germination and seem to have a low germination response to salinity. The lateral seeds, which are free from perianth envelopes, require light and cold pretreatment and seem to have a high sensitivity to salinity. Salicornia patula colonizes unstable transition zones between permanently flooded muds and perennial vegetation. These zones have a winter flooding period. The beginning of the flooding period and of the dry period may vary from year to year, depending on the erratic distribution of autumn and winter rains on the Mediterranean coast. So, in some years, a seedling population may be destroyed before reaching the stage of seed production. Seed dimorphism might cause germination to the more distributed in time, thus reducing the chances of extinction of a complete generation. In the present paper salinity and temperature response are described more precisely, are investigated, some aspects of the longevity of the two types of seed, and the adaptative significance of the corresponding dispersal and germination syndromes are discussed in more detail.     

Fruit and seed heteromorphism in the cold desert annual ephemeral Diptychocarpus strictus (Brassicaceae) and possible adaptive significance

Background and Aims

Diptychocarpus strictus is an annual ephemeral in the cold desert of northwest China that produces heteromorphic fruits and seeds. The primary aims of this study were to characterize the morphology and anatomy of fruits and seeds of this species and compare the role of fruit and seed hetermorphism in dispersal and germination.

Methods

Shape, size, mass and dispersal of siliques and seeds and the thickness of the mucilage layer on seeds were measured, and the anatomy of siliques and seeds, the role of seed mucilage in water absorption/dehydration, germination and adherence of seeds to soil particles, the role of pericarp of lower siliques in seed dormancy and seed after-ripening and germination phenology were studied using standard procedures.

Key Results

Plants produce dehiscent upper siliques with a thin pericarp containing seeds with large wings and a thick mucilage layer and indehiscent lower siliques with a thick pericarp containing nearly wingless seeds with a thin mucilage layer. The dispersal ability of seeds from the upper siliques was much greater than that of intact lower siliques. Mucilage increased the amount of water absorbed by seeds and decreased the rate of dehydration. Seeds with a thick mucilage layer adhered to soil particles much better than those with a thin mucilage layer or those from which mucilage had been removed. Fresh seeds were physiologically dormant and after-ripened during summer. Non-dormant seeds germinated to high percentages in light and in darkness. Germination of seeds from upper siliques is delayed until spring primarily by drought in summer and autumn, whereas the thick, indehiscent pericarp prevents germination for >1 year of seeds retained in lower siliques.

Conclusions

The life cycle of D. strictus is morphologically and physiologically adapted to the cold desert environment in time and space via a combination of characters associated with fruit and seed heteromorphism.     

Role of indehiscent pericarp in formation of soil seed bank in five cold desert Brassicaceae species

The dispersal and germination unit of some Brassicaceae species is the fruit, and we hypothesized that it could affect germination phenology and promote formation of a soil seed bank. We determined the effects of the indehiscent pericarp on germination and longevity of buried seeds of five Brassicaceae species native to cold deserts of central Asia. Germination phenology (seedling emergence) was monitored for intact dispersal units and isolated seeds of Chorispora sibirica, Goldbachia laevigata, Spirorrhynchus sabulosus, Tauscheria lasiocarpa (annuals), and Sterigmostemum fuhaiense (perennial) at natural temperatures in watered and non-watered (natural precipitation) soil. Intact dispersal units and isolated seeds were buried under natural conditions and exhumed at regular intervals for 35 months to monitor germination, viability and moisture content of isolated seeds, seeds in dispersal units, and seeds removed from dispersal units after burial. Isolated seeds of Goldbachia, Spirorrhynchus, and Tauscheria germinated only the first autumn and those of Chorispora and Sterigmostemum the first autumn and first spring, with higher germination percentages in all species in watered than in non-watered soil. A high percentage of seeds in buried dispersal units of Chorispora, Goldbachia, and Sterigmostemum was viable after 35 months, and seeds exhibited a 6-month dormancy cycle, being non-dormant only in autumn and spring. Seeds in buried dispersal units of Spirorrhynchus and Tauscheria germinated when exhumed in the first spring, but all non-germinated seeds were dead after 1 year. Thus, the presence of the pericarp allows Chorispora, Goldbachia, and Sterigmostemum to form a persistent seed bank but not Spirorrhynchus and Tauscheria.     

The impact of flooding and drought on seeds of Cnidium dubium, Gratiola officinalis, and Juncus atratus, three endangered perennial river corridor plants of Central European lowlands

The effects of flooding and drought on the post-flooding survival and germinability of seeds in the soil were investigated for three endangered river corridor angiosperms Cnidium dubium (Schkuhr) Thell., Gratiola officinalis L. and Juncus atratus Krocker. Freshly matured seeds were placed in small nylon bags and were buried in pots filled with soil collected from a wetland along the Lower Havel River. The pots were subjected to different winter/spring flooding treatments (30, 60, 120, and 180 days) and to drought for up to 3 years. Every year in spring, after drawdown, the seeds were exhumed, counted and tested for germination. Seed mortality of flooded seeds was low, except for C. dubium. Flooding did not affect germination of seeds of J. atratus. Flooding of seeds of C. dubium and G. officinalis had inconsistent, but no beneficial effects on germination of either. The year of harvest and the interannual variability of weather are found to be the main factors affecting germinability of C. dubium and G. officinalis in the course of the three consecutive years.The experiment showed that the capacity of the flooded seed bank to maintain germination ability and viability for extended periods of time could be directly related to the flooding duration of natural habitats of the three species. The three species are resilient to variable flooding. Seeds appeared desiccation tolerant, and single flooding events do not deplete their seed bank.     

The role of avian frugivores in germination of seeds of fleshy-fruited invasive alien plants

Many highly invasive plant species have fleshy fruits which are eaten by native frugivorous animals. These frugivores play an important role in long-distance seed dispersal, and may also affect germination success. The aim of this study was to determine whether generalist frugivores enhance or decrease seed germination of invasive alien species through pulp removal or seed coat abrasion, besides serving as dispersal agents. Fruits of four fleshy-fruited invasive alien plant species, namely Solanum mauritianum, Cinnamomum camphora, Lantana camara and Psidium guajava, were fed to three generalist avian frugivorous species, which have been observed feeding on these fruits in the wild. Seed retention time was recorded as this affects dispersal distance and the duration that seeds are exposed to the effects of the gut. Seeds removed from excreta, seeds from manually de-pulped fruit, and whole fruit were planted in soil trays housed in a greenhouse. Daily germination counts were done. Seed retention times differed significantly between bird species for all fruits, except those of C. camphora. However, all frugivores had a similar effect on the germination success of seeds of S. mauritianum, L. camara and P. guajava, showing that gut retention time was not important. Germination of seeds from manually de-pulped fruits did not differ from that of ingested seeds of all plant species, suggesting that seed coat abrasion was also not important. Pulp removal resulted in significantly higher germination rates, both in the two species with larger, multi-seeded fruit (S. mauritianum and P. guajava), and in the two species having single-seeded fruit with waxy exocarps (C. camphora and L. camara). Pulp removal also resulted in significantly earlier germination of L. camara and P. guajava seeds. Therefore, frugivores not only accelerate dispersal, but also greatly enhance seed germination of all fleshy-fruited invasive alien species in this study.