Christopher E. Mullins

GERMINATION ECOLOGY OF NEOTROPICAL PIONEERS: INTERACTING EFFECTS OF ENVIRONMENTAL CONDITIONS AND SEED SIZE

Germination provides many potentially unrecognized sources of variation in the regeneration niche. In this study we relate germination requirements and seed size for 16 species of pioneer trees to microclimatic conditions present in gaps in semi-deciduous rain forest in Panama. We found that, whereas increased duration of direct irradiance can be an effective indicator of the presence of a canopy gap across all scales of canopy openness, diel fluctuations in soil temperature effectively discriminate both understory sites and small gaps (25 m2) from larger gaps. Germination response was significantly related to seed size. Small-seeded species (seed mass 2 mg) germinated equally in light and darkness (wit...

Topographic position affects the water regime in a semideciduous tropical forest in Panamá

The effects of topographic position on water regime in a semideciduous tropical forest on Barro Colorado Island in Panamá were assessed by measuring soil matric potential using the filter paper technique and by using measured soil water release characteristics to convert a long-term (20 years) gravimetric water content data-set to matric potential. These were also compared against predictions from a simple water balance model. Soil matric potentials on slope sites were significantly higher than on plateau sites throughout the measurement interval and slopes experienced a shorter duration of drought during the annual dry-season. Measured values of matric potential agreed with those predicted from converting the gravimetric measurements using water release characteristics. Annual duration of drought predicted by the simple water balance model agreed with values determined from the converted long term water content data-set and was able to predict the annual duration of drought on plateau sites. On slope sites, the water balance systematically and significantly overestimated the duration of drought obtained from the water content data-set, suggesting that slope sites were supplied with water from upslope. Predictions of annual drought duration from sites with higher annual rainfall than Barro Colorado Island (BCI), suggest that while plateau sites on BCI experience a water regime consistent with annual rainfall, slopes experience a water regime more similar to that of forests with much higher rainfall. We conclude that such large variations in water regime over small spatial scales may play a role in maintaining high species richness through providing opportunities for niche specialisation and by buffering slopes against possible climate change.

Variable desiccation tolerance in Acer pseudoplatanus seeds in relation to developmental conditions: a case of phenotypic recalcitrance?

Nine seedlots of the widely planted southern and central European native tree species Acer pseudoplatanus L. were collected along a north-south gradient spanning 21° of latitude in Europe. We investigated how the heat sum during seed development influences seed maturity as assessed by physical, physiological and biochemical traits. Using principal component analysis we found predictable and consistent patterns in all traits, which correlated with heat sum. For example, compared with fruits from their native range (Italy and France, heat sum >3000°Cu2009d), fruits from the coldest location (Scotland; heat sum of 1873°Cu2009d) were shorter (c. 30 v. 42u2009mm), germinated over a narrower temperature range (5-20 v. 5-35°C) and had smaller embryos (28 v. > 70u2009mg) with a higher water content (c. 63 v. 48%), less negative solute potentials (c. -2.4 v. -4.1u2009MPa) and were more desiccation sensitive (critical water potential of -20.2 v. -55.4 to -60.7u2009MPa). The observed level of desiccation-tolerance for the French and Italian seedlots is more consistent with the intermediate category than the previous classification of A. pseudoplatanus as recalcitrant. Our results demonstrate that a lower heat sum causes fruits from northern Europe to be dispersed before maximum potential seed quality is achieved.

Germination Responses to Water Potential in Neotropical Pioneers Suggest Large-seeded Species Take More Risks

BACKGROUND AND AIMSnIn neotropical forests, very small-seeded pioneer species (<0.1 mg seed mass) recruit preferentially in small tree fall gaps and at gap edges, but large-seeded pioneers do not. Since water availability is related to gap size, these differences in microsite preference may reflect in part species-specific differences in germination at reduced water potentials.nnnMETHODSnFor 14 neotropical pioneer species, the hypothesis is tested that small-seeded species, with shallow initial rooting depths, reduce the risks associated with desiccation by germinating more slowly and at higher water potentials than large-seeded species.nnnKEY RESULTSnGermination occurred both more quickly and at lower water potentials with increasing seed mass. For example, Ochroma pyramidale (seed mass 5.5 mg) had a time to 50 % germination (T50) of 2.8 d and a median base potential for germination (psi(b50)) of -1.8 MPa while Clidemia quinquenervia (seed mass 0.017 mg) had a T50 of 17.6 d and psi(b50) of -1.1 MPa.nnnCONCLUSIONSnThese data suggest that small-seeded species germinate only in comparatively moist microsites, such as small canopy gaps, which may reduce the risk of drought-induced mortality. Conversely, large-seeded species are able to germinate in the drier environment of large gaps, where they benefit by enhanced seedling growth in a high irradiance environment. The positive association of seed size and canopy gap size for optimal seedling establishment is maintained by differential germination responses to soil water availability coupled with the scaling of radicle growth rate and seed size, which collectively confer greater drought tolerance on large-seeded species.

Effects of topographic position, leaf litter and seed size on seedling demography in a semi-deciduous tropical forest in Panama´

This study examined whether topography-induced gradients in water potential and leaf litter depth contribute to species coexistence in tropical forests through species-specific effects on seedling emergence and mortality. Seedling emergence and mortality were followed for a period of 12 months in 36 (1 × 2 m) plots on Barro Colorado Island (BCI), Panamá. Plots with and without litter were distributed on slope and plateau sites in three catchments. In the absence of manipulations, the lower litter depth on slopes resulted in approximately four times as many emergent seedlings than on plateau sites. However, litter depth had little effect on seedling community composition. By the end of the first dry-season, post-emergence, there were no significant differences in surviving seedling numbers between any treatments. There were differences in the emergent seedling community between slope and plateau sites within the same catchment as well as differences in composition between catchments, suggesting that both niche partitioning and dispersal limitation might play a role in structuring seedling community composition. During the wet-season seedling mortality was highest on slope sites although this pattern was reversed during the dry-season. In both seasons mortality was higher for small-seeded species. These results demonstrate that gradients in water potential related to topography impact on patterns of seedling emergence and mortality although processes in the first year after emergence may be insufficient to explain observed habitat preferences of adult plants.

Leaching of genetically modified Pseudomonas fluorescens through intact soil microcosms: Influence of soil type

SummaryThe leaching of a genetically modified Pseudomonas fluorescens through soil was investigated using intact (undisturbed) soil microcosms over a 2-month period. The microcosms comprised large cylindrical cores of three contrasting soil types (a loamy sand, a sandy loam, and a clay loam) supporting a grass/clover sward. Late log-phase cells of Pseudomonas fluorescens containing lux genes encoding for bioluminescence were applied to the surface of the soil cores. Eighteen hours after application of the inocula, the microcosms were subjected to simulated rain events (9 mm per event) at 3-day intervals and leachates were analysed for the concentration of genetically modified bacteria. The lux-modified pseudomonads were detected immediately in leachate from the clay looam with a steady decline in the concentration of cells with time. Leaching of pseudomonads from the sandy loam and loamy sand only occurred over a few rain events and total recoveries from the leachate were lower than from the clay loam. Leaching patterns are discussed in relation to differences in structure of topsoil and subsoil, which determine the pathways of water flow, and to the matric potential at inoculation, which determines the pore-size classes into which cells were first introduced.

Allometric relationships between seed mass and seedling characteristics reveal trade-offs for neotropical gap-dependent species

A seed size–seed number trade-off exists because smaller seeds are produced in greater number but have a lower probability of establishment. This reduced establishment success of smaller-seeded species may be determined by biophysical constraints imposed by scaling rules. Root and shoot diameter, root growth extension rate (RGER) and shoot length at death for dark-grown seedlings are predicted to scale with the cube root of seed embryo and endosperm mass (m). We confirmed this expectation for ten neotropical gap-dependent tree species with an embryo and endosperm dry mass >1xa0mg. However, for nine smaller seeded species (mxa0<xa01xa0mg) with photoblastic germination, root and shoot diameters were larger than expected, and consequently, RGER was slower than expected. The maximum shoot thrust of seedlings from seeds with masses ≥1xa0mg was comparable to the estimated force required to displace overlying litter, supporting the hypothesis that photoblastic behaviour only occurs in seeds with insufficient shoot thrust to displace overlying leaves. Using the model soil water, energy and transpiration to predict soil drying in small and large gaps, we showed that: (1) gaps that receive a significant amount of direct sunlight will dry more quickly than small gaps that do not, (2) compared to the wet-season, soil that is already dry at depth (i.e. the dry-season) will dry faster after rainfall (this drying would most likely kill seedlings from small seeds) and (3) even during the wet-season, dry periods of a few days in large gaps can kill shallow-rooted seedlings. We conclude that the smaller the seed, the more vulnerable its seedling would be to both covering by litter and soil drying because it can only emerge from shallow depths and has a slow RGER. Consequently, we suggest that these allometrically related factors contribute to the reduced establishment success of smaller-seeded species that underpins the seed size–seed number trade-off.

Burial and secondary dispersal of small seeds in a tropical forest

Secondary dispersal is an important stage in the life cycle of tree species, determining the fate of a high proportion of all seeds. For small-seeded species both physical and biological processes may influence the secondary fate of seeds, however the relative importance of these processes is not well known. Seeds of the pioneer tree species Cecropia insignis (seed mass 0.5mg), Trema micrantha (2.5mg) and Apeiba aspera (14.2mg) and five types of artificial seed were sown in understorey, treefall-gap and large-gap sites on Barro Colorado Island, Panama, during the wet season of 2005. Sowing areas were excavated after periods up to 26 d and cores divided into depths of 0-5, 5-10, 10-20and20-50mmtoallowhigh-resolutionestimationoftherateandamountofburialanddisplacementofseeds. Over 26 d, 2.8% of artificial seeds were buried to a mean depth of 10.5mm below the soil surface and 43.9% of unburied seeds displaced laterally >5cm. Significantly more (87.9% and 80.9%) seeds of Cecropia insignis and Trema micrantha were displaced than artificial seeds of similar mass, size and density. A generalised linear model suggested thatburialmostlyoccurredwithin15d,whiledisplacementoccurredcontinuouslyupto26d.Thedominantcauseof displacementandburialwasprobablyrainfall,whileseedremovalbyantsmayalsohavecontributedtodisplacement.