Sphagnum fallax is generally found closer to the water table than S. nemoreum along the hummock—hollow gradient of an Adirondack bog. However, at low water contents S. fallax unexpectedly fixes carbon at higher rates than S. nemoreum, as we have previously reported. We here compare water and carbon balances for these species to resolve this puzzling contrast. S. nemoreum maintains a more favorable water balance by virtue of its greater water—holding capacity, lower rates of water loss from moist plants, and apparently more effective capillary conduction of water. The consequence is a significantly higher tissue water content in the field compared to adjacent patches of S. fallax for a wide range of dates and heights above the water table. We developed a simulation model, PEATMOSS, to evaluate the relative importance of contrasting physiological relationships and water balances of the two species. The model integrates physiological and field data to estimate 24—h carbon balance for these mosses in adjacent patches at two heights above the water table under different conditions. PEATMOSS estimated a greater carbon gain for S. fallax at the low site, where it is a clear dominant over S. nemoreum, and a markedly more favorable carbon balance for S. nemoreum at the hummock site, where S. nemoreum is common but S. fallax is infrequent. Morphological features of S. nemoreum which effect its more favorable water balance apparently confer an advantage over S. fallax when water availability is limited.
The vertical distributions of nine Sphagnum species and four associated mosses in two bogs (Adirondack Mountains, New York) were analyzed for interspecific and interbog differences. Based on 4300 sample points, Sphagnum mean heights ranged from 12 to 43 cm above the water table and the sequence of species was similar in both bogs (S. cuspidatum and S. majus) < (S. fallax and S. angustifolium) < (S. magellanicum, S. rubellum, and S. russowii) < (S. fuscum and S. nemoreum). However, at Bloomingdale Bog, a relatively dry mire with many well-developed hummocks and narrow hollows, mosses had significantly greater means (by 4-12 cm) than at Raybrook Bog, a relatively wet mire with wide Sphagnum carpets and fewer hummocks. South- and east-facing hummock slopes tended to be gentler than the north and west slopes, but the mosses’ mean heights did not vary with aspect. The abundance of hummocks at Bloomingdale seems responsible for not only the greater mean heights, but also for broader ranges and greater vertical overlap with other species. However, the ability of each species to dominate a certain array of height classes did not differ between bogs. Within each bog, hummock species tended to have larger vertical ranges than hollow species, implying a lesser ability of hollow mosses to tolerate the full range of conditions along the hummock-hollow gradient.
The Eurasian watermilfoil (Myriophyllum spicatum L.) has partially replaced wild celery (Vallisneria americana Michx.) as a community dominant in the littoral zones of lakes of Madison, Wisconsin. The two species have very different growth forms, with that of M. spicatum corresponding more closely to the optimal growth form simulated by the macrophyte production model WEED. The objective of this research was to investigate the mechanisms by which Vallisneria could compensate for its nonoptimal growth form and coexist with Myriophyllum.A quantification of midsummer growth form for the two species at a rooting depth of 80-90 cm showed that M. spicatum had 68% of its shoot biomass within 30 cm of the surface, whereas V. americana had 62% of its leaf biomass within 30 cm of the bottom. Vallisneria had a light extinction coefficient ranging from 0.013 to 0.019 m·g, much higher than the value (ca. 0.006 m·g) for M. spicatum. This indicates less effective penetration of light to lower leaves of V. americana. Half-saturation constants describing the light-dependence of carbon uptake in "shade" and "sun" tissues ranged from 60-197 microeinsteins·m·s for V. americana, and 164-365 μeinsteins·m·s for M. spicatum. The optimum temperature for photosynthesis was 33.6°C for M. spicatum and 32.6°C for V. americana, but Myriophyllum was nearly twice as effective at carbon uptake at 10°C. Integration of all of the above features with WEED showed that, for midsummer conditions, V. americana more than compensated for apparently disadvantageous morphological features by its greater physiological adaptability to low light regimes. Coupled with the temperature-dependence of photosynthesis, it appears that V. americana is favored by midsummer conditions, whereas M. spicatum is at an advantage at other times.
Sphagnum fallax and S. nemoreum differ in their vertical distribution above the water table on hummocks in Bloomingdale Bog (Adironkack Mountains, New York), and both exhibit decreasing water content with increasing height above the water table. We tested the prediction that S. nemoreum, a species better developed on hummock tops, would photosynthesize at greater rates when relatively dry than would S. fallax, which is more abundant in hollows and on hummock bases. Net photosynthesis declined sharply at tissue water contents (fresh—/dry—mass ratio) below 7.2 for S. fallax and 8.9 for S. nemoreum collected in spring. These values fell to 5.1 and 7.4, respectively, for plants collected in late summer. Thus both species exhibit adaptive seasonal acclimation, as reflected by the maintenance of relatively high net photosynthetic rates to lower water contents after a dry summer. Neither species showed significant response differences when collected from different heights above the water table. In direct contradiction to our hypothesis, Sphagnum fallax fixed carbon at greater rates than S. nemoreum at low water contents for both spring and summer collections. This physiological contrast appears inconsistent with observed field distribution patterns for these species.
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