Impacts of sedimentation and nitrogen enrichment on wetland plant community development

Many factors influence which plant species are found in a particular wetland. The species pool is composed of the species present in the seed bank and species able to disperse into the wetland, and many abiotic and biotic factors interact to influence a species performance and abundance in the plant community. Anthropogenic activities produce specific stressors on wetland systems that alter these abiotic and biotic interactions, potentially altering species composition. We simulated three common wetland hydrogeomorphic (HGM) subclasses in a greenhouse to examine the effects of two stressors-sedimentation and nitrogen (N) enrichment-on the performance of 8 species grown in artificial communities. Species establishment, height, biomass, and foliar N and P concentrations were measured to explore species responses to stressors and competition, as well as the potential impacts of changes in species composition on ecosystem processes. Species were affected differently by sedimentation and N enrichment, and there were differences in overall community sensitivity to stressors between wetland subclasses. Sedimentation generally reduced seedling establishment, while N enrichment produced variable effects on height and biomass. Interspecific competition had little effect on establishment but significantly reduced most species biomass. Sedimentation generally lowered community biomass, diversity, and richness, while enrichment increased community biomass. Establishment, biomass, and foliar nutrient concentrations significantly differed between many species, suggesting that shifts in species composition may impact ecosystem processes such as nutrient cycling and carbon storage. Phalaris arundinacea, an aggressive clonal graminoid, universally dominated all wetland subclasses. This dominance across a range of environmental conditions (sedimentation, fertility, and hydrology) has important implications for both restoration and predicting the impacts of human activities on species composition. Our results suggest that, in regions where P. arundinacea is common, restoration projects that establish communities from seeds and human activities that cause vegetation removal are likely to become dominated by P. arundinacea.     

Growth of Phragmites australis and Phalaris arundinacea in constructed wetlands for wastewater treatment in the Czech Republic

Common reed (Phragmites australis) and reed canarygrass (Phalaris arundinacea) are two most commonly used plant species in constructed wetlands for wastewater treatment in the Czech Republic. Growth characteristics of both plants (biomass, stem count, and length) have been measured in 13 horizontal sub-surface flow constructed wetlands since 1992. The results revealed that while Phalaris usually reaches its maximum biomass as early as during the second growing season, Phragmites usually reaches its maximum only after three to four growing seasons. The maximum biomass of both species varies widely among systems and the highest measured values (5070 g m⁻² for Phragmites and 1900 g m⁻² for Phalaris) are similar to those found in eutrophic natural stands. The shoot count of Phragmites decreases after the second growing season while length and weight of individual shoots increases over time due to self-thinning process. Number of Phalaris shoots is the highest during the second season and then the shoot count remains about the same. Also the shoot length remains steady over years of constructed wetland operation.     

Establishment of Carex stricta Lam. seedlings in experimental wetlands with implications for restoration

The loss of Carex dominated meadows due to agricultural drainage in the previously glaciated midcontinent of North America has been extensive. The lack of natural Carex recruitment in wetland restorations and the failures of revegetation attempts underscore the need for information on the establishment requirements of wetland sedges. In this study, seedlings of Carex stricta Lam. were planted in three experimental wetlands in east-central Minnesota, USA to investigate the biotic and abiotic environmental limitations to establishment. Seedlings were planted along an elevational water depth gradient to assess the effects of water depth and water level fluctuation on seedling survival and growth. A different water level fluctuation regime was assigned to each of the experimental wetlands to assess seedling tolerance for seasonal water level changes. The effects of seedling planting density and the presence or absence of non-sedge colonizers on seedling survival and growth were also studied. The experiment was followed for three growing seasons. The results of this study indicate that C. stricta seedlings were sensitive to the timing and duration of inundation during the first growing season. Once established, plants tolerated a broad range of seasonal drying and flooding conditions. Seedling and juvenile growth was slowed by non-sedge colonizers during the first two growing seasons, but by the third growing season, C. stricta was able to out-grow all annual and perennial weeds, except the aggressive perennial, Phalaris arundinacea L. The rapid growth of C. stricta plants, once established, indicates that the use of seedlings is a successful method for (re) introducing this tussock sedge into wetland restorations under a variety of environmental conditions. Comparison with other studies performed under similar conditions suggests that planting of seedlings is a more appropriate method of establishing this species than the use of transplanted rhizomes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Comparison of created and natural freshwater emergent wetlands in Connecticut (USA)

Five three- to four-year old created palustrine/emergent wetland sites were compared with five nearby natural wetlands of comparable size and type. Hydrologic, soil and vegetation data were compiled over a nearly two-year period (1988-90). Created sites, which were located along major highways, exhibited more open water, greater water depth, and greater fluctuation in water depth than natural wetlands. Typical wetland soils exhibiting mottling and organic accumulation were wanting in created sites as compared with natural sites. Typha latifolia (common cattail) was the characteristic emergent vegetation at created sites, whereas a more diverse mosaic of emergent wetland species was often associated with Typha at the natural sites. Species richness was slightly higher in created (22–45) vs. natural (20–39) wetlands, but the mean difference (33 vs. 30) was not significant. Nearly half (44%) of the 54 wetland taxa found at the various study sites were more frequently recorded at created than natural wetlands. The presence of mycorrhizae in roots of Typha angustifolia (narrow-leaved cattail) and Phragmites australis (common reed) was greater at created than natural wetlands, which may be related to differential nutrient availability. Wildlife use at all sites ranged from occasional to rare, with more sightings of different species in the natural (39) than created (29) wetlands. The presence of P. australis and introduced Lythrum salicaria (purple loosestrife) may pose a threat to future species richness at the created sites. One created site has permanent flow-through hydrology, and its vegetation and wildlife somewhat mimic a natural wetland; however, the presence of P. australis and its potential spread pose an uncertain future for this site. This study suggests the possibility of creating small palustrine/emergent wetlands having certain functions associated with natural wetlands, such as flood water storage, sediment accretion and wildlife habitat. It is premature to evaluate fully the outcome of these wetland creation efforts. A decade or more is needed, emphasizing the importance of long term monitoring and the need to establish demonstration areas.     

Disturbance, Herbivory, and Propagule Dispersal Control Dominance of an Invasive Grass

Despite the dramatic changes invasive plants cause to ecosystems and communities, factors that control dominance of invasive species after establishment in a community are poorly understood. Most active management relies on catastrophic disturbances of invasive-dominated communities to increase richness and diversity of plant communities. This study examines the importance of propagule dispersal and deer herbivory on continued dominance of Phalaris arundinacea after a non-catastrophic, short-term disturbance to monotypic stands of this invasive grass. The disturbance caused no change in P. arundinacea cover among treatments during any year of the study and, thus, simulates disturbance intensity more likely to be encountered in unmanaged settings. Despite the small disturbance, the combinations of disturbance + seeding and disturbance + seeding + deer exclusion caused greater species richness than controls even three years after disturbance. Increased invasion of P. arundinacea stands caused few effects on the dominant, as P. arundinacea biomass was unaffected after the first year. Selective herbivory by deer of species other than P. arundinacea increased the effects of disturbance and seeding, and aided in continued dominance of the grass. The tolerance of P. arundinacea for direct anthropogenic effects, including poor water quality and hydroperiod fluctuations, and indirect effects, such as increased herbivory by historically high deer populations, indicates the complexity of determining persistence of invasive species.     

Fish community structure,spatial distribution and feeding ecology in a beaver pond

Synopsis The fish community of a small (2.7 ha) Ontario beaver pond was analyzed relative to predictions based on its small size, shallow depth, brief existence and isolation from more permanent water bodies. The predictions were: (1) species richness will be lower than that of more permanent water bodies in the area, (2) fish will be mainly of small body size, (3) species will be randomly distributed across habitats, and (4) there will be a high degree of diet overlap between species and age classes. The first and second predictions were supported. The pond consisted of 10 resident species in 1985, and at least seven in 1988. Species richness was below the average of 13.2 found in four lakes in the vicinity, but greater than the 3.1 predicted by a species-area curve for non-acidified lakes in Ontario. All species except pumpkinseed, yellow perch and brown bullheads were small-bodied with short life spans and high population turnover rates, and few fish above 100 mm were present. Predictions 3 and 4 were not supported. Habitat occupation was nonrandom, and high diet separation occurred, particularly in August when food limitation was evident. While beaver ponds lack the range of habitats and the diversity of species of lake environments, low prey density and high fish density nevertheless appear to foster resource partitioning.     

Influence of hydrological fluctuations on the growth and nutrient dynamics ofPhalaris arundinacea L. in a riparian environment

Seasonal changes in aboveground and belowground tissues ofPhalaris arundinacea L. were studied in a population colonizing an ancient meander of the Garonne river (France) submitted to important fluctuations of the permanent water table. Waterlogged conditions in spring stopped the growth of rhizomes and promoted the translocation of nutrient to the shoots. The early senescence of plants after flowering could be related to the withdrawal of the water table. It was characterized by a distribution of nutrients in belowground tissues and a release in litter and soil. Aerated conditions in late summer permitted the growth of belowground tissues. At this time a partition of resources between aboveground and belowground biomass of a new generation of plants was observed. Rising water and decreasing temperatures in winter induced the death of aboveground parts. Reconstitution of nutrient stocks in rhizomes and losses by leaching then occured. Beside a very high primary production this strategy confers toPhalaris arundinacea a great interest in different uses, especially in the removal of nutrients from water in riparian zones as in artificial sites.     

Seasonal variation in seed bank composition and its interaction with nutrient enrichment in the Everglades wetlands

In the Florida Everglades, nutrient enrichment from agricultural outflow and the change in hydrology have collectively contributed to the expansion of cattails (Typha spp.). To assess the effectiveness of prescribed fire in controlling cattails and to predict vegetation dynamics after the fire, it is important to understand the seasonal variation of the soil seed bank and how the seed bank is affected by nutrient enrichment and fire. This paper investigates the effects of season, nutrient enrichment, and fire on soil seed bank species composition, richness, and density along a nutrient gradient in Water Conservation Area 2A (WCA 2A) of the Florida Everglades. Species richness was significantly affected by nutrient enrichment and season but not their interaction. Total seed density, however, was significantly affected by the interaction between nutrient enrichment and season. Yet, at species level, the relationship between seed density, nutrient enrichment and season varied. The highest seed density of cattail occurred in summer at highly enriched sites, but that of sawgrass occurred in fall regardless of enrichment; the seed density of water lily was very low regardless of season and nutrient enrichment, and the highest Amarathus seed density occurred at highly enriched sites year round. Moreover, germination timing differed greatly among species. While cattail seeds had a short incubation period and started to germinate 2–3 days after initiation of the germination assay, sawgrass seeds generally started to germinate 4 weeks later. Further, both the prescribed summer fire at the highly enriched site and the natural winter fire at the moderately enriched site reduced the seed density of cattail but not of sawgrass. Our results suggest that fire application for vegetation recovery in WCA 2A would benefit from explicitly considering seasonal dynamics of the seed bank.     

Habitat preferences of cattail species and hybrids (Typha spp.) in eastern Canada

Interspecific hybridization is widespread in plants and is an important evolutionary process. Hybrids may be fitter than their parental species, at least under some environmental conditions, and this may lead to partitioning of taxa by habitat. In eastern Canada, two cattail species (Typha latifolia and Typha angustifolia) and their hybrid (Typha x glauca) are known to have become increasingly widespread in recent decades, although their habitat preferences and patterns of co-existence at the local scale are not well known. We quantified the occurrence of these three taxa in three different habitat types (high traffic, low traffic, and ‘natural’) at 40 different sites along a sampling route of approximately 2000 km in eastern Canada. There were no significant overall associations between habitat type and taxon, although intraspecific comparisons among sites showed that the hybrid was most likely to grow in high traffic (highly disturbed) sites. In addition, pairwise comparisons revealed significant independence of T. latifolia and T. angustifolia, although the hybrid was equally likely to co-exist with either of its parental species. The presence of the three taxa in several habitats, including highly disturbed roadside areas, is consistent with their increasingly invasive tendencies.     

A comparison of Phragmites australisin freshwater and brackish marsh environments in North America

This paper compares the available North Americanliterature and data concerning several ecologicalfactors affecting Phragmites australisin inlandfreshwater, tidal fresh, and tidal brackish marshsystems. We compare aboveground productivity, plantspecies diversity, and sediment biogeochemistry; andwe summarize Phragmiteseffects on faunalpopulations in these habitats. These data suggest thatPhragmitesaboveground biomass is higher thanthat of other plant species occurring in the samemarsh system. Available data do not indicate anysignificant difference in the aboveground Phragmitesbiomass between marsh types, nor doesthere appear to be an effect of salinity on height.However, Phragmitesstem density wassignificantly lower in inland non-tidal freshwatermarshes than in tidal marshes, whether fresh orbrackish. Studies of the effects of Phragmiteson plant species richness suggest that Phragmitesdominated sites have lower diversity.Furthermore, Phragmiteseradication infreshwater sites increased plant diversity in allcases. Phragmitesdominated communities appearto have different patterns of nitrogen cyclingcompared to adjacent plant communities. Abovegroundstanding stocks of nitrogen (N) were found to behigher in Phragmitessites compared to thosewithout Phragmites. Porewater ammonium(NH₄ ⁺) did not differ among plant covertypes in the freshwater tidal wetlands, but inbrackish marshes NH₄ ⁺was much higher inSpartinaspp. than in neighboring Phragmitesstands. Faunal uses of Phragmitesdominated sites in North America were found to vary bytaxa and in some cases equaled or exceeded use ofother robust emergent plant communities. In light ofthese findings, we make recommendations for futureresearch.     

Molecular genetic data reveal hybridization between Typha angustifolia and Typha latifolia across a broad spatial scale in eastern North America

A recent increase in the abundance of cattails (Typha spp.) in North American wetlands has been anecdotally linked with hybridization between Typha latifolia and Typha angustifolia. In this study, we used molecular genetic markers (microsatellites) to investigate whether the hybrid lineage (Typha × glauca) is restricted to The Great Lakes region, or exists across a much broader spatial scale. We also investigated the possibility of backcrossing and genetic introgression in natural populations. Parental species could be distinguished from one another based on the distribution of alleles at six microsatellite loci. Species identification based on genetic data corresponded well with species identifications based on leaf width, a key morphological trait that can distinguish the two parental species. We found that hybrids occur in Ontario, Quebec, New Brunswick, and Nova Scotia, but we did not detect hybrids in Maine. F₁s are more abundant than backcrossed or intercrossed hybrids, although we also found evidence of backcrossing, particularly in Ontario. This indicates that hybrids are fertile, and are therefore potential conduits of gene flow between the parental species. Further work is needed to determine whether T. × glauca is particularly successful in the Great Lakes region relative to other areas in which the two parental species co-exist, and to assess whether introgression may lead to increased invasiveness in the species complex.     

Growth and biomass turnover ofHydrocharis dubia L. cultured under different nutrient conditions

Growth of a floating-leaved plant,Hydrocharis dubia L., was examined under varying nutrient conditions between 0.3 and 30 mgN l⁻¹ total inorganic nitrogen.H. dubia plants cultured under the most nutrient-rich condition showed the highest maximum ramet density (736 m⁻²), the highest maximum biomass (80.4 g dry weight m⁻²), and the highest total net production (185 g dry weight m⁻² in 82 days). Plants under nutrient-poor conditions had a relatively large proportion of root biomass and a small proportion of leaves with a long life span. Compared with other floating-leaved and terrestrial plants, the maximum biomass ofH. dubia was relatively small. This, and the rapid biomass turnover, was related to the short life span of leaves (13.2–18.7 days) and large biomass distribution to leaves.     

Managing vegetation in surface-flow wastewater-treatment wetlands for optimal treatment performance

Constructed wetlands that mimic natural marshes have been used as low-cost alternatives to conventional secondary or tertiary wastewater treatment in the U.S. for at least 30 years. However, the general level of understanding of internal treatment processes and their relation to vegetation and habitat quality has not grown in proportion to the popularity of these systems. We have studied internal processes in surface-flow constructed wastewater-treatment wetlands throughout the southwestern U.S. since 1990. At any given time, the water quality, hydraulics, water temperature, soil chemistry, available oxygen, microbial communities, macroinvertebrates, and vegetation each greatly affect the treatment capabilities of the wetland. Inside the wetland, each of these components plays a functional role and the treatment outcome depends upon how the various components interact. Vegetation plays a uniquely important role in water treatment due to the large number of functions it supports, particularly with regard to nitrogen transformations. However, it has been our experience that vegetation management is critical for achieving and sustaining optimal treatment function. Effective water treatment function and good wildlife quality within a surface-flow constructed wetland depend upon the health and sustainability of the vegetation. We suggest that an effective tool to manage and sustain healthy vegetation is the use of hummocks, which are shallow emergent plant beds within the wetland, positioned perpendicular to the water flow path and surrounded by water sufficiently deep to limit further emergent vegetation expansion. In this paper, we describe the use of a hummock configuration, in conjunction with seasonal water level fluctuations, to manage the vegetation and maintain the treatment function of wastewater-treatment wetlands on a sustainable basis.     

Morphology of the gallery system around the nest and gallery development under experimental conditions in the termiteMacrotermes bellicosus (Smeathman)

Summary In the Ivory Coast (Côte d'Ivoire) we analysed the structures and the lay-out of the underground gallery system of the fungus-growing termiteMacrotermes bellicosus. We found a radial pattern of principal and lateral galleries, as well as caves, depressions and vertical passages reaching a depth of 1–2 m. According to the results of our investigations the gallery networks built byM. bellicosus are unique among all known termite species;M. bellicosus constructs the densest (3.1 m/m²) and most superficial (2–6 cm under the surface) gallery networks among all comparable termite species investigated.Under experimental conditions involving an observation arena with a glass bottom, we were able to observe the dynamics of the radial development of the exploratory passages. We also observed the manner and rapidity of the termites' exploration of new ground, as well as the role played by the discovery of food in the design of the gallery system.We were able to show that, when food was scarce, the termites gained access to the arena much more rapidly, built a much denser gallery system and came up to the surface more frequently.     

Clonal spread of invasive Ludwigia hexapetala and L. grandiflora in freshwater wetlands of California

Ludwigia hexapetala and L. grandiflora are recent, aggressive invaders of freshwater wetlands in California. To assess the relative role of sexual versus clonal reproduction in invasive spread, we used AFLP markers to genotype 794 ramets of L. hexapetala sampled from 27 populations in three watersheds of California, and 150 ramets of L. grandiflora from five populations in a fourth watershed. We then used two analytical methods, similarity thresholds and character incompatibilities, to distinguish genotypic variation within genets (clones) from variation between genets. Our analyses revealed extremely limited genotypic and genet variation in invasive L. hexapetala and L. grandiflora within California. Within L. hexapetala, 95% of the ramets analyzed represented a single genet. The genet was the only one detected in 20 populations. The remaining seven populations contained two to nine genets. Within L. grandiflora, all ramets were of only one genotype. Thus, invasive spread within and between populations, and across watersheds, appears to be almost exclusively clonal and brought about by the dispersal of vegetative propagules. The extremely low seedling recruitment indicates that management should target vegetative dispersal and growth.     

O-acetylserine (thiol) lyase activity in Phragmites and Typha plants under cadmium and NaCl stress conditions and the involvement of ABA in the stress response

The roles of O-acetylserine (thiol) lyase (OASTL, EC 4.2.99.8) and abscisic (ABA) acid in stress responses to NaCl and cadmium treatments were investigated in Typha latifolia L. and Phragmites australis (Cav.) Trin. ex Steudel plants. OASTL activity increased under stress (25-300 microM Cd, 100mM NaCl, 1 microM ABA) in both Typha and Phragmites mainly in roots, contributing substantially to satisfy the higher demand of cysteine for adaptation and protection. The earliest significant responses in intact roots were recorded after 12-24 h of Cd treatments, but different levels of stimulation were also observed after 3 and 7 days of exposure. The OASTL activity responses of Phragmites to salinity (100mM NaCl) were higher than those of Typha. Cysteine synthesis in Typha is much higher than in Phragmites, which supports the efficiency of the thiol-metabolism-based protection shown in Typha. Exogenous ABA increased OASTL activity in both species. Cd treatments led to increased ABA levels in roots. Phragmites showed higher ABA levels compared to Typha. The increase of ABA content indicates the involvement of this phytohormone in early stress responses, while the stimulation of OASTL following the ABA application suggests that ABA has a role in an OASTL activation pathway.     

Geographic locality and host identity shape fungal endophyte communities in cupressaceous trees

Understanding how fungal endophyte communities differ in abundance, diversity, taxonomic composition, and host affinity over the geographic ranges of their hosts is key to understanding the ecology and evolutionary context of endophyte–plant associations. We examined endophytes associated with healthy photosynthetic tissues of three closely related tree species in the Cupressaceae (Coniferales): two native species within their natural ranges [Juniperus virginiana in a mesic semideciduous forest, North Carolina (NC); Cupressus arizonica, under xeric conditions, Arizona (AZ)], and a non-native species planted in each site (Platycladus orientalis). Endophytes were recovered from 229 of 960 tissue segments and represented at least 35 species of Ascomycota. Isolation frequency was more than threefold greater for plants in NC than in AZ, and was 2.5 (AZ) to four (NC) times greater for non-native Platycladus than for Cupressus or Juniperus. Analyses of ITS rDNA for 109 representative isolates showed that endophyte diversity was more than twofold greater in NC than in AZ, and that endophytes recovered in AZ were more likely to be host-generalists relative to those in NC. Different endophyte genera dominated the assemblages of each host species/locality combination, but in both localities, Platycladus harboured less diverse and more cosmopolitan endophytes than did either native host. Parsimony and Bayesian analyses for four classes of Ascomycota (Dothideomycetes, Sordariomycetes, Pezizomycetes, Eurotiomycetes) based on LSU rDNA data (ca 1.2 kb) showed that well-supported clades of endophytes frequently contained representatives of a single locality or host species, underscoring the importance of both geography and host identity in shaping a given plant's endophyte community. Together, our data show that not only do the abundance, diversity, and taxonomic composition of endophyte communities differ as a function of host identity and locality, but that host affinities of those communities are variable as well.     

Effects of Cu,Pb and Zn on two Potamogeton species grown under field conditions

Two common macrophyte species, Potamogeton perfoliatus L. and Potamogeton pectinatus L. were grown for 12 weeks at shallow depths in sediments contaminated with 1250 or 2500 g Pb or Cu and/or Zn (gDW sediment)^-1. Control experiments were run at background levels of 4, 13, and 38 g Pb, Cu and Zn (gDW sediment)^-1, respectively. Effects of heavy metals on biomass production and metal uptake and distribution in plants are presented in relation to total amount and plant-available fraction of metals in the sediment.All three studied metals gave reduced biomass production, and the toxicity of the metals decreased in the order Zn>Cu>Pb. The root/shoot biomass ratio increased for P. pectinatus, but decreased for P. perfoliatus with metal treatment. The content of any single metal was higher in shoots than in roots of plants grown on sediments not contaminated with that specific metal, but addition of that metal increased the proportion in roots. The uptake by plants of any of the heavy metals increased with increased metal addition. The magnitude of the plant-available fraction of metals of untreated sediment was Zn>Cu>Pb, and increased in contaminated sediments. Addition of Cu decreased both the plant-available fraction and the total concentration of Zn in the sediment, while increased the uptake of Zn by the plants. The opposite was found for Cu when Zn was added. P. pectinatus accumulated about twice as much Cu as P. perfoliatus. On the other hand, the concentration of Pb was higher in P. perfoliatus than in P. pectinatus, and was negligible in P. pectinatus when cultivated in untreated sediments.     

Comparison of heavy metal accumulation in a natural wetland and constructed wetlands receiving acid mine drainage

Metal accumulations in sediments and plants of constructed and natural wetlands were compared in two wetlands constructed by the Tennessee Valley Authority (TVA) for the treatment of acid mine drainage and a natural wetland. Load rates and removal efficiencies of most metals were generally greater in the constructed wetlands than in the natural wetland. There were similar sediment and plant metal concentrations between one constructed wetland and the natural wetland and greater metal concentrations in the sediments and plants in the other constructed wetland compared to the natural wetland. Data indicate that Mn, Zn, Cu, Ni, B, and Cr are being accumulated in the plants at all three wetlands, although accumulation of metals by these plants accounts for only a small percentage of the removal of the annual metal load supplied to each wetland.     

Response of heart rate and cloacal ventilation in the bimodally respiring freshwater turtle, Rheodytes leukops, to experimental changes in aquatic PO2

Changes in heart rate (f _H) and cloacal ventilation frequency (f _C) were investigated in the Fitzroy turtle, Rheodytes leukops, under normoxic (17.85 kPa) and hypoxic (3.79 kPa) conditions at 25°C. Given R. leukops’ high reliance on aquatic respiration via the cloacal bursae, the objective of this study was to examine the effect of varying aquatic PO₂ levels upon the expression of a bradycardia in a freely diving, bimodally respiring turtle. In normoxia, mean diving f _H and f _C for R. leukops remained constant with increasing submergence length, indicating that a bradycardia failed to develop during extended dives of up to 3 days. Alternatively, exposure to aquatic hypoxia resulted in the expression of a bradycardia as recorded by a decreasing mean diving f _H with increasing dive duration. The observed bradycardia is attributed to a hypoxic-induced metabolic depression, possibly facilitated by a concurrent decrease in f _C. Results suggest that R. leukops alters its strategy from aquatic O₂ extraction via cloacal respiration in normoxia to O₂ conservation when exposed to aquatic hypoxia for the purpose of extending dive duration. Upon surfacing, a significant tachycardia was observed for R. leukops regardless of aquatic PO₂, presumably functioning to rapidly equilibrate blood and tissue gas tensions with alveolar gas to reduce surfacing duration.