Rhizome dynamics and resource storage in Phragmites australis

Seasonal changes in rhizome concentrations of total nonstructural carbohydrates (TNC), water soluble carbohydrates (WSC), and mineral nutrients (N, P and K) were monitored in two Phragmites australis stands in southern Sweden. Rhizome biomass, rhizome length per unit ground area, and specific weight (weight/ length ratio) of the rhizomes were monitored in one of the stands.Rhizome biomass decreased during spring, increased during summer and decreased during winter. However, changes in spring and summer were small (< 500 g DW m-2) compared to the mean rhizome biomass (approximately 3000 g DW m^–2). Winter losses were larger, approximately 1000 g DW m-2, and to a substantial extent involved structural biomass, indicating rhizome mortality. Seasonal changes in rhizome length per unit ground area revealed a rhizome mortality of about 30% during the winter period, and also indicated that an intensive period of formation of new rhizomes occurred in June.Rhizome concentrations of TNC and WSC decreased during the spring, when carbohydrates were translocated to support shoot growth. However, rhizome standing stock of TNC remained large (> 1000 g m^–2). Concentrations and standing stocks of mineral nutrients decreased during spring/ early summer and increased during summer/ fall. Only N, however, showed a pattern consistent with a spring depletion caused by translocation to shoots. This pattern indicates sufficient root uptake of P and K to support spring growth, and supports other evidence that N is generally the limiting mineral nutrient for Phragmites.The biomass data, as well as increased rhizome specific weight and TNC concentrations, clearly suggests that reloading of rhizomes with energy reserves starts in June, not towards the end of the growing season as has been suggested previously. This resource allocation strategy of Phragmites has consequences for vegetation management.Our data indicate that carbohydrate reserves are much larger than needed to support spring growth. We propose that large stores are needed to ensure establishment of spring shoots when deep water or stochastic environmental events, such as high rhizome mortality in winter or loss of spring shoots due to late season frost, increase the demand for reserves.     

Life history and reproductive potential of the agarophyte Gelidium robustum in California

The reproductive potential of the tetrasporangial phase of Gelidium robustum was studied for 16 months at two sites off Santa Barbara, California. In all samples tetrasporangial thalli were always more abundant than gametangial ones. Tetratrasporangial sori were present throughout the duration of the study but relative fecundity was highest [300–400 sori g^–1 (w. wt)] in spring/summer samples of consecutive years, as a result of increasing numbers both of tetrasporangial branchlets per plant and of sori per branchlet. On the other hand, laboratory experiments showed that tetraspore release per sorus was highest (150–250 spores sorus^–1 d^–1) in winter. Inferring from these field and laboratory data plants released up to ± 34 000 tetraspores g^–1 (w. wt) d^–1 in the spring/summer of the second study year. Tetraspore germination, under defined culture conditions, also showed a marked seasonality increasing sharply from less than 10% in winter up to almost 60% in spring/summer, thus coinciding with the period of maximal spore output per plant. These results suggest that although relatively high numbers of tetraspores may be released by G. robustum plants all year round these might not always have the potential to germinate and recruit.     

Seasonal variation in the physical properties of agar and biomass of Gracilaria sp. (chorda type) from Tosa Bay,southern Japan

Plants of Gracilaria sp.(chorda type), which grow along the coast of Uranouchi Inlet in Tosa Bay, southern Japan, showed the highest biomass in the summer (26 °C to 31 °C) and spring season (15.1 °C to 24.9 °C). Maximum biomass was 6952 g m^–2 in July, but gradually decreased in the autumn (30.5 °C in September to 20 °C in November) and winter (19.5 °C in December to 14.9 °C in February). Variation in yields and gel strength of the agars, were shown to depend on the time in the season. After alkali treatment (5% NaOH, 2 h) at three different temperatures (70, 80, and 90 °C), the agars showed gel strengths essentially that of commercial grade agars, with the best gel obtained at 80 °C. Maximum gel strength (1455 g cm^–2 of 1.5% agar gel) occurred in winter when the biomass and agar yield were low. Minimum gel strength was in spring. Gel strength was inversely correlated with agar yield, but was positively correlated with apparent viscosity. Maximum viscosity was 40 cP. in December. Gelling temperatures, pH of 1.5% agar gel, and moisture content in agars showed little variation.     

Relationship between a high density of sika deer and productivity of the short-grass (<Emphasis Type="Italic">Zoysia japonica</Emphasis>) community: a case study on Kinkazan Island,northern Japan

An extremely high-density (ca. 800 deer km^–2) wild sika deer population uses a short-grass community dominated by Zoysia japonica on Kinkazan Island in northeastern Japan. To explain why the density of wild deer is quite high on the Zoysia community, (1) we quantified the seasonal productivity of the Zoysia community, (2) we compared food availabilities among the plant communities, and (3) we described the habitat selection by the deer in different seasons. Food availability was greater on the Zoysia community than in the forest understory from spring to fall. The productivity of the Zoysia community was high enough to support the actual high density of the deer (814 deer km^–2) in summer. However, the productivity markedly decreased in winter, when the deer density decreased to less than half (358 deer km^–2) of the summer value. In contrast, the deer density of the adjacent forests was highest in winter (154 deer km^–2) and lowest in spring (19 deer km^–2). These results suggest that the deer using the Zoysia community in summer left and were absorbed into the adjacent forest in winter. If such an adjacent community were absent, many deer would not survive, and consequently the deer density on the Zoysia community in summer would not be so high. This intercommunity movement is particularly important for the deer using a plant community like the Zoysia community, which is highly productive but has a small standing biomass.     

Food habits of sambar <Emphasis Type="Italic">Cervus unicolor</Emphasis> at the Horton Plains National Park,Sri Lanka

The sambar Cervus unicolor has a large population in southern Asia. In Sri Lanka, its habitat use is most common among the four cervid species. We carried out an analysis of the food habits of sambar in the Horton Plains National Park from January to November 1991 by using the quantitative fecal analysis method. Analyses of sambar feces collected at the three habitats (two grassland communities and a forest) in the Horton Plains National Park found that the sambar diet largely consisted of graminoid leaves; 45–78% in grasslands and 43–60% in the forest. The analyses also found that the proportion of graminoid leaves in early summer (52–78%) was greater than it was in winter (43–49%). In spring and summer, different varieties of graminoids were consumed by sambar, whereas dicotyledonous plants were consumed in winter. The fecal nitrogen level did not fall below the critical level of their nutrition (1.1%), indicating that sambar food quality in the Horton Plains was sufficient. A large proportion of Pennisetum spp., exotic grasses (10–39%) were found in the fecal compositions collected in the Pennisetum community. This could be because of the high biomass and high nitrogen concentration (approximately 2.4%) of Pennisetum and its low-growing nature. From the present study, we found that in order for sambar, a native ungulate, not to increase in number, the growth of exotic plants such as Pennisetum needs to be controlled.     

Annual and seasonal changes in fine root biomass of a Quercus ilex L. forest

The biomass, production and mortality of fine roots (roots with diameter <2.5 mm) were studied in a typical Mediterranean holm oak (Quercus ilex L.) forest in NE Spain using the minirhizotron methodology. A total of 1212 roots were monitored between June of 1994 and March of 1997. Mean annual fine root biomass in the holm oak forest of Prades was 71±8 g m^–2 yr^–1. Mean annual production for the period analysed was 260+11 g m^–2 yr^–1. Mortality was similar to production, with a mean value of 253±3 g m^–2 yr^–1. Seasonal fine root biomass presented a cyclic behaviour, with higher values in autumn and winter and lower in spring and summer. Production was highest in winter, and mortality in spring. In summer, production and mortality values were the lowest for the year. Production values in autumn and spring were very similar. The vertical distribution of fine root biomass decreased with increasing depth except for the top 10–20 cm, where values were lower than immediately below. Production and mortality values were similar between 10 and 50 cm depth. In the 0–10 cm and the 50–60 cm depth intervals, both production and mortality were lower.     

Life history, aspects of reproductive biology and production of Corophium orientale (Crustacea: Amphipoda) in Monolimni lagoon (Evros Delta, North Aegean Sea)

Monthly samples of Corophium orientale were collected during February 1998–February 1999 in both parts of Monolimni lagoon (0.3 < Sal. < 6 psu, 2 < Temp. < 28.5 °C). Corophium orientaleshowed a semiannual life cycle. In the southern part, breeding occurred from early spring to mid autumn having three peaks, in early spring, early summer and early autumn, and three cohorts were produced, a spring cohort, a summer cohort and an overwintering one. In the shallower innermost northern part of Monolimni lagoon, breeding peaked in mid-spring and, also, in early autumn, and two discrete cohorts, a spring and an overwintering one, were produced. A lack of large individuals, attributed to a temporary size – selective predation by migrating shorebirds, was observed during summer. That lack possibly contributed to the hiatus in reproduction. Photoperiod seemed to be a more important cue than temperature in the initiation of the reproductive cycle, while extremely low salinities (<1 psu) did not deter breeding. Females attained maturity at a smaller size in late spring and, especially, in summer than in early spring at lower temperatures. Brood size was a function of females body size. Females, matured during summer at high temperatures, showed the lowest brood size, body length ratio. Mean brood size was small (12–13 early embryos) and embryo loss during development high (54%), possibly due to a negative effect of low salinities. A 1:1 sex ratio existed in the small body length classes, but females preponderated in the large ones. The spring cohort in the southern part, which developed at moderate temperatures, showed the highest growth rate (40 m day^–1). Secondary production of C. orientalecalculated by Hyness method gave a mean annual density of 4562.5 and 9327.6 ind. m^–2, a mean annual crop (B) of 1.03 and 2.67 g DW m^–2, an annual production (P) of 6.91 and 22.54 g DW m^–2 and a P/B ratio of 6.7 and 8.4 in the northern and southern part of the lagoon, respectively.     

Summer biomass of a population of Iridaea cordata (Gigartinaceae,Rhodophyta) from Antarctica

Results of a seasonal study on biomass in an infralittoral population of Iridaea cordata from Terra Nova Bay (Ross Sea, Antarctica) are reported. Thalli were collected during the IX Italian Antarctic Expedition (austral summer 1993–94). The population studied is that living at depths of 4 to 6 m, where the highest density of plants occurred. The highest value of biomass (wet weight 3440 g m^–2) was found at the beginning of summer. In that period 72.5% of biomass was from 128 specimens belonging to weight classes 8 (>16 to 32 g) to 10 (>64 g), corresponding to 13.4% of the population in numbers. Small (<1 g) and medium (1 to 8 g) specimens provided the remaining biomass of 5% and 22.5%, respectively. During the month of January, the number of heavy specimens decreased. At the end of that month biomass reached a minimum of 2225 g m^–2. In February the biomass increased to 3169 g m ^–2, 72% of which was from 120 specimens belonging to weight classes 7 (>8 to 16 g) to 9 (>32 to 64 g), which numerically represented 18.5% of the population. Data showed that biomass depended mainly on the presence of large heavy specimens, even though they were always few in number. Moreover, the occurrence of such large thalli at the beginning of summer suggests that Iridaea cordata continues to grow during the long antarctic winter.     

Metazooplankton dynamics in the Salton Sea, California, 1997–1999

The dynamics of metazooplankton populations were studied over 3 years at the saline (43 g l^–1) Salton Sea, California's largest lake. Total abundance was highest in summer following late winter/early spring phytoplankton blooms. At this time, metazooplankton consisted mostly of the copepod, Apocyclops dengizicus, and the rotifer, Brachionus rotundiformis. In August or September, severe crashes in the metazooplankton populations occurred each year in mid-lake due to strong wind events which increased mixing and caused low oxygen and high sulfide concentrations throughout the water column. Larvae of the polychaete worm, Neanthes succinea and the barnacle, Balanus amphitrite were present mostly in late winter and spring. Their scarcity in summer is due in part to persistent anoxic bottom conditions that decrease adult populations and in part to predation by tilapia, an omnivorous fish that has become abundant in the lake since the 1960s. Two Synchaeta species, rotifers not previously reported from the Sea, were abundant in winter and spring and predation on these may have permitted the copepod to persist at low levels through the winter. There were two major changes in metazooplankton dynamics since 1954–1956 in addition to the appearance of the two synchaetid rotifers in the fauna. First, there are now much lower densities of barnacle and polychaete larvae in the fall, probably due to the invasion of the zooplanktivorous fish, tilapia. Second the precipitous crashes now seen in metazooplankton densities, especially the copepod, in late summer-early fall did not occur in the 1950s possibly because fall overturn events did not result in such high sulfide levels.     

Field energetics of a large carnivorous lizard,Varanus rosenbergi

Summary Field metabolic rates (FMRs; CO₂ production) and water influx rates of Varanus rosenbergi were measured seasonally by means of doubly-labelled water (³H₂ ¹⁸O). Metabolic rates and water influexes were highest in summer (0.18 ml CO₂ g^–1 h^–1 and 17.9 ml H₂O kg^–1 day^–1) and lowest in winter (0.04 ml CO₂ g^–1 h^–1 and 6.5 ml H₂O kg^–1 day^–1). FMRs and water fluxes were intermediate in spring. It is estimated that a 1-kg V. rosenbergi would need to consume 4.7 kg of prey each year to remain in energy balance and that the energy commitment to egg production requires a 40% increase in the food acquisition of a gravid female in late spring and early summer. It is calculated that water influxes in summer are totally provided from food but in other seasons, particularly winter, pulmocutaneous water exchange and/or drinking are significant avenues of water intake.     

Phytoplankton communities in channel catfish ponds

Seasonal changes in the quantity and quality of phytoplankton were studied in six channel catfish culture ponds. Chlorophyll a concentrations were generally highest in the summer (averaging >200 g 1^–1) but the highest individual chlorophyll a value recorded (910 g 1^–1) occurred in the winter during a bloom of Dictyosphaerium pulchellum. On the average, green algae (Chlorophyta) and euglenoids (Euglenophyta) represented relatively constant proportions of the phytoplankton community seasonally (about 35 and 10%, respectively). In the summer and fall, blue-green algae (Cyanophyta) became abundant. Diatoms were relatively abundant at all times and constituted the majority of the community in the winter and spring.     

Field Note: Comparative Efficacy of A Woody Evapotranspiration Landfill Cover Following the Removal of Aboveground Biomass

Woody vegetation cultivated for moisture management on evapotranspiration (ET) landfill covers could potentially serve a secondary function as a biomass crop. However, research is required to evaluate the extent to which trees could be harvested from ET covers without significantly impacting their moisture management function. This study investigated the drainage through a six-year-old, primarily poplar/cottonwood ET test cover for a period of one year following the harvest of all woody biomass exceeding a height of 30 cm above ground surface. Results were compared to previously reported drainage observed during the years leading up to the coppice event. In the first year following coppice, the ET cover was found to be 93% effective at redirecting moisture during the spring/summer season, and 95% effective during the subsequent fall/winter season. This was slightly lower than the 95% and 100% efficacy observed in the spring/summer and fall/winter seasons, respectively, during the final measured year prior to coppice. However, the post-coppice efficacy was higher than the efficacy observed during the first three years following establishment of the cover. While additional longer-term studies are recommended, this project demonstrated that woody ET covers could potentially produce harvestable biomass while still effectively managing aerial moisture.     

The predation impact of juvenile herring Clupea harengusand sprat Sprattus sprattus on estuarine zooplankton

The consumption of estuarine copepods by juvenile herring and sprat during estuarine residency was estimated using fish biomass data and daily rations calculated from two models of feeding in fish: a bioenergetic model and a gastric evacuation model. The bioenergetic model predicted daily rations that were, on average, three times higher than those estimated by a model based on field records of stomach contents. The biomass of herring and sprat in the estuary was negatively correlated with the daily ration suggesting that the clupeid fish populations were resource-limited. Copepod production decreased towards the winter and peaked in spring and summer. The relative importance of predation changed seasonally in function of the migration pattern of herring and sprat. In the spring and the summer, in situ production ofcopepod biomass was higher than the in situ consumption by fish. During the fall and the winter, consumption exceeded production. This suggests that top–down control exerted by marine pelagic fish may be an important force structuring estuarine copepod populations.     

Botanical and chemical composition of rumen contents of Sika deer on Mt Goyo,northern Japan

Botanical and chemical compositions of the rumen contents of 58 Sika deer on Mt Goyo, northern Japan, collected from summer of 1988 to spring of 1989, were analyzed.Sasa nipponica, a dwarf bamboo, was important in summer (35.0%) and winter (61.4%), but it decreased to 5.6% and was replaced by browse leaves in fall, and to 28.0% and was replaced by dead leaves, twigs and bark of woody plants in winter. Crude protein was 20–25% lower in the washed fraction than in the gross fraction. It was highest (16.2%) in summer and lowest (8.6%) in winter. High protein content in summer and fall foods was attributed to forb and browse leaves. Seasonal fluctuation in protein content in the foods of these Sika deer was greater than red deer on Rhum, Scotland and smaller than wapiti in Yellowstone National Park, Wyoming, USA. Climatically, winter severity of Mt Goyo is intermediate between the two localities, which seems to explain the seasonal fluctuation of the protein level. Crude fiber wasca 33% in the ‘washed’ fraction, and did not change seasonally.     

Estimating seasonal and annual carbon inputs,and root decomposition rates in a temperate pasture following field 14C pulse-labelling

Using a ¹⁴C pulse-labelling technique, we studied the seasonal changes in assimilation and partitioning of photoassimilated C in the plant–root–soil components of a temperate pasture. Pasture and soil samples were taken after 4-h, and 35-day chase periods, to examine these seasonal ¹⁴C fluxes. Total C and ¹⁴C were determined in the shoot, root and soil system. The amounts of C translocated annually to roots and soil were also estimated from the seasonal ¹⁴C distribution and pasture growth. The in situ field decomposition of newly formed roots during different seasons, also using ¹⁴C-labelling, was studied for one year in undisturbed rhizosphere soil. The ¹⁴C-labelled roots were sampled five times and decomposition rates were calculated assuming first-order decomposition.Annual pasture production at the site was 16 020 kg DM ha^–1, and pasture growth varied with season being highest (75–79 kg ha^–1 d^–1) in spring and lowest (18–20 kg ha^–1 d^–1) in winter. The above- and below-ground partitioning of ¹⁴C also varied with the season. The respiratory ¹⁴C–CO₂ losses, calculated as the difference between the total amounts of ¹⁴C recovered in the soil-plant system at 4 h and 35 days, were high (66–70%) during the summer, autumn and winter season, and low (37–39%) during the spring and late-spring season. Pasture plants partitioned more C below-ground during spring compared with summer, autumn and winter seasons. Overall, at this high fertility dairy pasture site, 18 220 kg C/ha was respired, 6490 kg remained above-ground in the shoot, and 6820 kg was translocated to roots and 1320 kg to soil. Root decomposition rate constant (k) differed widely with the season and were the highest for the autumn roots. The half-life was highest (111 days) for autumn roots and lowest (64 days) for spring roots. About one-third of the root label measured in the spring season disappeared in the first 5 weeks after the initial 35 Day of allocation period. The late spring, summer, late summer and winter roots had intermediate half-lives (88–94 days). These results indicate that seasonal changes in root growth and decomposition should be accounted for to give a better quantification of root turnover.     

Root biomass,root distribution and the fine-root growth dynamics of Quercus coccifera L. in the garrigue of southern France

Quercus coccifera L., the characteristic scrub oak of the garrigue, covers more than 100,000 ha in southern France alone. Precipitation in this area averages 900 mm/year and summer rains are not rare. A total belowground biomass of 7.2 kg/m², including rhizomes and lignotubers, was harvested. Roots were concentrated in the uppermost 50 cm of the soil. It was hypothesized that low winter temperatures inhibit active fine-root growth. This hypothesis was tested by means of fine-root extractions of soil samples from 0–50 cm depth from November 1987 to June 1988. Although the fine-root analysis could not be extended into late summer and fall, the data supported the hypothesis. Ratios of live/dead fine roots reached their minimum at 0.2–0.3 from December to April. They increased to 1.0–1.2 during late spring and early summer. Initiation of fine-root growth in early April was synchronous with bud break. Starch contents of roots, rhizomes, and lignotubers fluctuated from 4.3% in January to 8.3% in April. The starch stored in belowground organs of Q. coccifera in a closed canopy stand amounted to about 500 g/m² in April. This amount declined to 400 g with bud burst and fine-root growth initiation.     

BIOMASS,PRODUCTION, AND LITTERFALL IN THE COASTAL SAGE SCRUB OF SOUTHERN CALIFORNIA

A 22-year-old stand of coastal sage scrub in the coastal mountains of southern California had a peak standing aboveground biomass of 1,417 g/m², determined by dimension analysis. Annual aboveground net primary production was 255 g/m²/yr, determined by monthly twig harvests of dominant species and the clipping production of subordinate species. The stand was codominated by two drought-deciduous species, Salvia leucophylla and Artemisia californica, which together comprised 81% of the biomass. Annual litterfall was measured at 194 g/m²/yr. These biomass, production, and litterfall values are less than those measured in most evergreen chaparral communities in California. Seasonally, the two dominant shrubs began aboveground production in the winter soon after the first rains and continued growth for six months until early summer. A massive leaf fall occurred in May–June as the summer drought began, but twig and inflorescence production for both species continued at a high rate into the summer months. Salvia leucophylla had two shoot types: 1) an early spring canopy shoot that elongated rapidly, produced the inflorescence, and died in mid-summer; and 2) a short side shoot produced in late spring with small dense leaves that were retained during the summer drought and early winter. Artemisia californica produced a single cohort of twigs in the early spring, most of which carried inflorescences by late summer.     

Seasonal variations in the encapsulation of eggs of the encyrtid parasitoid Metaphycus stanleyi by the pyriform scale, Protopulvinaria pyriformis

Encapsulation of eggs of the introduced parasitoid Metaphycus stanleyi Compere (Hymenoptera: Encyrtidae) by the pyriform scale, Protopulvinaria pyriformis (Cockerell) (Homoptera: Coccidae) under both greenhouse and field conditions, at Bet Dagan, was found to occur almost all year round (1986–1988). However, encapsulation rates varied considerably during the different seasons and were correlated with the ambient temperatures. The rates of efficient encapsulation (i) in scales infesting Hedera helix and Schefflera arboricola under greenhouse conditions, were lowest during December to May (6–17%) and highest during July to September (78–100%); (ii) in scales infesting avocado in the orchard, were lowest during October to May (0–11%) and highest during June to August (54–57%). Under greenhouse conditions, encapsulation rates did not differ in scales grown on H. helix and on S. arboricola, but were significantly lower in scales grown on avocado. Encapsulation by scales infesting S. arboricola was more frequent at temperatures ranging from 20–28 °C, than from 7–23 °C.The increased resistance by encapsulation of P. pyriformis to successful parasitization by M. stanleyi during the summer, may account for the inability of the parasitoid to prevent the autumn and winter outbreaks of the pest.     

Seasonal changes in bacterial and phytoplankton biomass in a subantarctic coastal area (Kerguelen Islands)

The seasonal variations of bacterial and phytoplanktonic biomass were studied during several pluri-annual surveys in the subantarctic Morbihan Bay (Kerguelen Islands, 49 ° 20 S; 70 ° 10 E). Large interannual variation was observed. Phytoplanktonic biomass showed moderate values during winter and autumn. They increased sharply in spring, reaching a maximum value of about 1 mg C l^–1 corresponding to an important depletion of nutrients. A second phytoplanktonic bloom of similar amplitude occurred in late summer. During algal blooms which were roughly associated with optimal values of solar irradiation for the first one and with the highest temperatures for the second one, phytoplanktonic material represented near 100% of particulate and living carbon. Bacteria showed maximal abundance (0.2 to 0.7 mg C l^–1) during summer or autumn. Their relative abundance, which represented less than 1% of the living biomass in spring and summer, can reach more than 95% in autumn and winter.     

Seasonal flooding regimes influence survival,nitrogen fixation,and the partitioning of nitrogen and biomass in Alnus incana ssp. rugosa

Alteration of natural flooding regimes can expose lowlands to waterlogged soil conditions during any month of the year. The seasonality of flooding may have profound effects on the carbon and nitrogen budgets of N-fixing alders (Alnus spp.), and in turn, may impact the C and N economy of extensive alder-dominated, wetland ecosystems, including those dominated by speckled alder (Alnus incana ssp. rugosa). To better understand this process, two-year-old, nodulated seedlings of speckled alder were subjected to late spring (May 10 – July 10), summer (July 10 – September 8), and fall (September 8 – November 8) flooding treatments. Alders were root-flooded outdoors in tanks containing an N-free nutrient solution and compared with unflooded alders at the experimental site. Flooding arrested N fixation, photosynthesis, and growth of alders without recovery in all flooding treatments for the remainder of the growing season. Late spring and summer flooding resulted in complete mortality of alders while all seedlings survived flooding in the fall. Fall flooding increased foliar N resorption by 140% over unflooded seedlings. Eighty-seven percent of the total N fixed and 89% of biomass accumulation for the entire growing season occurred in unflooded alders after July 10. In unflooded alders, nitrogen fixation rates per unit mass declined by 63% for nodules, 28% for leaves, and 48% for whole seedlings during the fall, while total N fixed per plant in the fall was similar to that fixed in the summer. The majority of newly fixed N in unflooded alders was allocated to leaves before September 8 and to roots/nodules combined after September 8. In unflooded plants, the greatest proportion of new biomass was partitioned to leaves before July 10, to stems between July 10 and September 8, and equally to stems and roots/nodules after September 8. Fall-flooded alders did not increase root or nodule biomass. Proportional allocation of plant resources were such that the ratio of N fixed to seedling growth of unflooded alders decreased by 19% during summer before rebounding by 6% in fall. Seasonality of flooding alters seedling survival, growth, and resource allocation, and may be a critical determinant of speckled alder recruitment and occurrence in wetlands.