Responses of macroinvertebrate communities to long‐term flow variability in a Sonoran Desert stream

Current global models predict a hotter and drier climate in the southwestern United States with anticipated increases in drought frequency and severity coupled with changes in flash flood regimes. Such changes would likely have important ecological consequences, particularly for stream and riparian ecosystems already subject to frequent hydrologic disturbance. This study assessed the potential response of aquatic macroinvertebrates to interannual variation in hydrology in a spatially intermittent desert stream (Sycamore Creek, AZ). We compiled data on the recovery of macroinvertebrate communities following spring floods, with successional sequences captured 11 times over a 16‐year period (1983–1999). This period encompassed a transition from perennial to intermittent flow in this system, and included a record drought in 1989–1990. Results show that while the size of floods initiating sequences had little explanatory power, changes in macroinvertebrate community structure during postflood succession were closely associated with antecedent flooding and drought. Year‐to‐year differences in benthic communities integrated taxon‐specific responses to antecedent disturbance, including differential resistance to channel drying, use of hyporheic refugia, and variable rates of recovery once stream flow resumed. The long‐term consequences of drying on community structure were only evident during later stages of postflood succession, illustrating an interaction between flood and drought recovery processes in this system. Our observations highlight the potential for predicted climate changes in this region to have marked and long‐lasting consequences for benthic communities in desert streams.     

溪流生态系统潜流带生态学研究概述

最近一、二十年,对潜流带研究的逐渐开展已使人们充分认识到潜流带在溪流生态系统结构、功能和生态过程中的重要意义。潜流带是溪流地表水和地下水相互作用的界面,占据着在地表水、河道之下的可渗透的沉积缓冲带、侧向的河岸带和地下水之间的中心位置。有关潜流带重要性的近期研究进展增进了我们对溪流生态学的理解,大大扩展了水生生物生境的物理空间以及生物相互作用和生产力存在的区域。潜流带生境包含多样化的、丰度很高的动物区系,其常常控制着溪流生物生产力和生态过程。潜流带对溪流生态系统的潜在重要性来自于生物学和生物地化活动。潜流带在溪流有机物的分解和水质净化等生态过程中发挥着重要的作用。潜流带是溪流集水区内各种环境特征的综合表征体,它能够反映周围景观的环境条件及其变化,集水区内的自然或人为干扰,都会影响到潜流带的水文循环、理化特征、生境分布、营养结构和基本生态过程。因此,对溪流生态系统的综合管理必须包括潜流带。最后对潜流带生态学研究现存的主要问题及未来研究展望提出了几点看法。     

Flood effects on invertebrates, sediments and particulate organic matter in the hyporheic zone of a gravel-bed stream

1. We investigated the effects of a flood on the fauna and physical habitat of the hyporheic zone of the Kye Burn, a fourth order gravel‐bed stream in New Zealand. 2. Freeze core hyporheic samples (to 50 cm depth) and benthic samples (to 10 cm) were taken, along with measurements of vertical hydrological gradient, before, 2 days after and 1 month after the flood (estimated return period: 1.5 years, estimated Q_max = 10.4 m³ s^?1). 3. The composition of the hyporheos differed over the three sampling occasions with fewer taxa collected immediately postflood than preflood. The equitability of the community was higher on both postflood occasions, consistent with the reduced densities of two abundant taxa (Leptophlebiidae and Copepoda). 4. Total invertebrate abundance was lower on the postflood occasions than preflood in both benthic (0–10 cm) and hyporheic (10–50 cm) sediments. Several taxa, including asellotan isopods and amphipods, recovered within 1 month of the event. Hyporheic densities of larval Hydora and nematodes did not differ among the three sampling occasions, but the water mite Pseudotryssaturus was more abundant 1 month after the flood than preflood. There was no evidence of vertical movements (to 50 cm) by any taxa in response to the flood. 5. The proportion of fine sediments (<1 mm) in the subsurface sediments (10–50 cm) increased over the three sampling occasions and median particle size declined, but sediment porosity did not change. More particulate organic matter was found in the sediments after the flood. 6. Our study provides little evidence that the hyporheic zone (to 50 cm) acted as a significant refuge during the flood event, although movements to or recolonisation from sediments deeper than 50 cm could explain the recovery of many crustacean and mite taxa within 1 month.     

Nitrogen processing in the hyporheic zone of a pastoral stream

The distribution of nitrogen-transforming processes, and factors controlling their rates, were determined within the hyporheic zone of a lowland stream draining agricultural land. In the field, physicochemical parameters were measured along a 10m-long hyporheic flow line between downwelling and upwelling zones. Sediment cores were retrieved from the stream bed surface, and from 20, 40 and 60cm deep in each zone, and in the laboratory, water from the corresponding depth was percolated through each core at the natural flow rate. Concentrations of nitrogen species and oxygen were measured before and after flow through each core. Denitrification was measured using a ¹⁵N-nitrate tracer. Shallow and downwelling zone samples were clearly distinct from deeper and upwelling zone samples in terms of physicochemical conditions, microbial processes and factors controlling nitrogen processing. Denitrification was highest in surface and downwelling zone cores, despite high oxygen levels, probably due to high pore-water nitrate concentrations in these cores and isolation of the denitrifying bacteria from oxygen in the bulk water by the hyporheic biofilms. Denitrification was limited by oxygen inhibition in the downwelling group, and by nitrate availability in the upwelling group. Strong evidence indicated that dissimilatory nitrate reduction to ammonium, occurred in almost all cores, and outcompeted denitrification for nitrate. In contrast, nitrification was undetectable in all but two cores, probably because of intense competition for oxygen. Field patterns and lab experiments indicated that the hyporheic zone at this moderately N-rich site is a strong sink for nitrate, fitting current theories that predict where hyporheic zones are nitrate sinks or nitrate sources.     

Seasonal reversals of upland-riparian diversity gradients in the Sonoran Desert

Flood disturbance and water resource availability vary sharply over time and space along arid‐region rivers and can interact in complex fashion to shape diversity patterns. Plant diversity showed spatial patterning along a topogradient from the floodplain of the San Pedro River (Arizona, USA) to the arid upland, but the patterns shifted temporally as the suite of limiting factors changed. During two of three sampling times, spatial diversity patterns were shaped primarily by gradients of water availability, the regional limiting factor. In the summer dry season, microscale diversity (species richness per 1 m²) and mesoscale diversity (cumulative species and functional types in 20, 1‐m² plots) of herbaceous plants decreased along the topogradient from floodplain to upland, reflecting the greater water availability on the low surfaces. During a summer wet season with moderate rains and flooding, diversity increased in all hydrogeomorphic zones (floodplain, terrace, upland), but the spatial pattern along the topogradient persisted. Following a very wet winter, patterns along the topogradient reversed: scour from large floods limited diversity on the floodplain and competitive exclusion limited the diversity on undisturbed river terrace, while abundant rains allowed for high microscale diversity in the upland. Disturbance and resource availability thus interacted to influence plant species diversity in a fashion consistent with the dynamic‐equilibrium model of species diversity. In contrast to the microscale patterns, mesoscale diversity of species and functional types remained high in the floodplain during all sampling times, with 58% more plant species and 90% more functional types sampled in low floodplain than arid upland for the year as a whole. Species with a wide range of moisture and temperature affinities were present in the floodplain, and seasonal turnover of species was high in this zone. The floodplain zone of a perennial to intermittent‐flow river thus had greater plant diversity than arid Sonoran Desert upland, as measured at temporal scales that capture seasonal variance in resource and disturbance pulses and at spatial scales that capture the environmental heterogeneity of floodplains. Although periodically limited by intense flood disturbance, diversity remains high in the floodplain because of the combination of moderate resource levels (groundwater, seasonal flood water) and persistent effects of flood disturbance (high spatial heterogeneity, absence of competitive exclusion), in concert with the same climatic factors that produce seasonally high diversity in the region (temporally variable pulses of rainfall).     

山地河流浅滩生境潜流层大型无脊椎动物群落拓殖

通过包埋人工基质法研究大型无脊椎动物在山地河流潜流层中的拓殖过程。结果表明:群落个体密度在7—29 d呈"J"型增长,在29 d后骤然降低,55 d后呈波动趋势;物种丰富度在1—29 d呈增加趋势,29 d后呈波动状态;群落的生物量总体呈增加趋势。群落的物种丰富度、密度和生物量在第29、71和83天时没有显著性差异(P0.05),Shannon-Wiener多样性指数和Pielou均匀度指数在第55、71、83天没有显著性差异(P0.05),综合不同拓殖时间段物种的主成分分析,表明潜流层大型无脊椎动物群落在55 d后趋于稳定。群落优势种为摇蚊(Camptochironomus sp.)、河蚬(Corbicula fluminea)、四节蜉(Baetis sp.)、动蜉(Cinygmina sp.)、纹石蛾(Hydropsyche sp.)和扁泥甲科的一种(Psephenidae)。滤食者和收集者在整个拓殖过程中均是优势功能摄食群。群落拓殖过程是一个群落自身恢复能力和外部环境影响相互作用的过程,拓殖初期潜流层的结构是影响着无脊椎动物迁入的主要因素,中期动物的生活史特征起主要作用,稳定期之后群落可能受到各因素的综合影响。     

Nitrogen dynamics in the hyporheic zone of a forested stream during a small storm, Hokkaido, Japan

Water and dissolved nitrogen flows through the hyporheic zone of a 3rd-order mountain stream in Hokkaido, northern Japan were measured during a small storm in August 1997. A network of wells was established to measure water table elevations and to collect water samples to analyze dissolved nitrogen concentrations. Hydraulic conductivity and the depth to bedrock were surveyed. We parameterized the groundwater flow model, MODFLOW, to quantify subsurface flows of both stream water and soil water through the hyporheic zone. MODFLOW simulations suggest that soil water inflow from the adjacent hill slope increased by 1.7-fold during a small storm. Dissolved organic nitrogen (DON) and ammonium (NH ₄ ⁺ ) in soil water from the hill slope were the dominant nitrogen inputs to the riparian zone. DON was consumed via mineralization to NH ₄ ⁺ in the hyporheic zone. NH ₄ ⁺ was the dominant nitrogen species in the subsurface, and showed a net release during both base and storm flow. Nitrate appeared to be lost to denitrification or immobilized by microorganisms and/or vegetation in the riparian zone. Our results indicated that the riparian and hyporheic system was a net source of NH ₄ ⁺ to the stream.     

A mixing model analysis of stream solute dynamics and the contribution of a hyporheic zone to ecosystem function*

1. We monitored streamwater and streambed sediment porewaters from White Clay Creek (WCC), SE Pennsylvania, for dissolved organic carbon (DOC), dissolved oxygen (DO) and conductivity to investigate organic matter processing within the hyporheic zone. Dissolved organic carbon and DO concentrations were higher in the streamwater than in the porewaters and, in many cases, concentrations continued to diminish with increasing depth into the streambed. 2. Hydrological exchange data demonstrated that the permeability of the stream bed declines with depth and constrains downwelling, effectively isolating porewaters >30 cm from streamwater. 3. End‐member mixing analysis (EMMA) based on conductivity documented a DOC source and DO sink in the hyporheic zone. We calculated hyporheic streambed DOC fluxes and respiration from the EMMA results and estimates of water flux. Based upon our calculations of biodegradable DOC entering the hyporheic zone, we estimate that DOC supports 39% of the hyporheic zone respiration, with the remaining 61% presumably being supported by entrained particulate organic carbon. Hyporheic respiration averaged 0.38 g C m^?2 d^?1, accounted for 41% of whole ecosystem respiration, and increased baseflow ecosystem efficiency from 46 to 59%. 4. Advective transport of labile organic molecules into the streambed concentrates microbial activity in near‐surface regions of the hyporheic zone. Steep gradients in biogeochemical activity could explain how a shallow and hydrologically constrained hyporheic zone can dramatically influence organic matter processing at the ecosystem scale.     

Bacterial community dynamics in the hyporheic zone of an intermittent stream

The dynamics of in situ bacterial communities in the hyporheic zone of an intermittent stream were described in high spatiotemporal detail. We assessed community dynamics in stream sediments and interstitial pore water over a two-year period using terminal-restriction fragment length polymorphism. Here, we show that sediments remained saturated despite months of drought and limited hydrologic connectivity. The intermittency of stream surface water affected interstitial pore water communities more than hyporheic sediment communities. Seasonal changes in bacterial community composition was significantly associated with water intermittency, phosphate concentrations, temperature, nitrate and dissolved organic carbon (DOC) concentrations. During periods of low- to no-surface water, communities changed from being rich in operational taxonomic units (OTUs) in isolated surface pools, to a few OTUs overall, including an overall decline in both common and rare taxa. Individual OTUs were compared between porewater and sediments. A total of 19% of identified OTUs existed in both porewater and sediment samples, suggesting that bacteria use hyporheic sediments as a type of refuge from dessication, transported through hydrologically connected pore spaces. Stream intermittency impacted bacterial diversity on rapid timescales (that is, within days), below-ground and in the hyporheic zone. Owing to the coupling of intermittent streams to the surrounding watershed, we stress the importance of understanding connectivity at the pore scale, consequences for below-ground and above-ground biodiversity and nutrient processing, and across both short- and long-time periods (that is, days to months to years).     

溪流潜流层大型无脊椎动物生态学研究进展

溪流潜流层是溪流表层水和地下水相互作用的群落交错区,生物多样性丰富,是溪流生态系统的重要组成部分.大型无脊椎动物位于潜流层食物网的顶层,直接影响着潜流层物质和能量动态,是河流健康潜在的指示生物,调节着潜流层的环境净化和生态缓冲功能,对溪流生态系统发挥着至关重要的作用.潜流层大型无脊椎动物类群按生活史划分为偶入动物、非典型潜流层动物和典型潜流层动物.潜流层孔隙大小、孔隙水流速、溶解氧、温度、可利用的食物源、渗透系数和水力停留时间是影响大型无脊椎动物在潜流层分布的主要因素.对于潜流层这样一个特殊的生态界面,针对不同的研究目的应该选择合适的取样和调查方法.潜流层大型无脊椎动物的生活史和生活史对策,在溪流生态系统物质循环和能量流动中作用的定量化分析,基于潜流层大型无脊椎动物的河流健康评价体系,以及潜流层作为“庇护地”对于大型无脊椎动物分布和进化的生态学意义,都值得进一步关注和深入研究.     

Seasonal and spatial distribution of ciliates in the sandy hyporheic zone of a lowland stream

The seasonal and spatial distribution of abundance and biomass as well as the taxonomic composition of ciliates inhabiting the sandy hyporheic zone of a lowland stream were studied. The mean abundances varied between 0 and 895 cells ml⁻¹ sediment, and the mean ciliate biomass ranged between 0 and 5.3 μg C ml⁻¹ sediment. Ciliate numbers and biomasses were greatest at the sediment surface and declined significantly with increasing sample depth. Abundance and biomass varied seasonally, with maximum values in late autumn and early winter and minimum values in early summer. The community was dominated by small representatives of the Hymenostomatia and Peritrichia. Ciliate community composition changed with depth from a very diverse community at the sediment surface to a less diverse one at greater sediment depths. Ciliate abundance and biomass were two orders of magnitude lower in the channel water than in the hyporheic zone. Although representatives of all sediment taxa could also be found in the channel water, the greatest concentrations of Peritrichia and Suctoria were in the hyporheic zone. The species of the sandy Ladberger Mühlenbach sediment were ubiquitous; there was no single ciliate fauna that proved to be typical for this kind of freshwater biotope.     

The hyporheic zone: Hydrogeological and geochemical study of a stream in the pyrenees mountains

The hydrogeological and geochemical characteristics of the hyporheic zone were studied for a year on a 30 m² site within a stream in the Pyrenees Mountains. This included the drawing of potentiometric maps, measuring hydraulic conductivity and porosity, and the monthly analysis of major chemical elements in water samples. Use of the combined data led to the clarification of certain aspects of the characteristics, functioning and development of the hyporheic zone. Even though the area studied lies in a zone of groundwater recharge, the direct vertical exchange between surface- and groundwater does not seem to be determinant in the potentiometric and chemical development of the area. However, the state of reserves, which can be reduced or reconstituted, seems to play an essential role in flow dynamics, and thus of the chemistry of this area. Flow dynamics in the hyporheic zone apparently are of the longitudinal-transversal type, related to the local stream morphology. The results show how difficult it is to define the hyporheic zone in a structural sense, even though the zone represents an entity that is clearly characterized by specific and identifiable dynamics.     

Reproductive Consequences of Clonal Growth in Stenocereus eruca, a Rare Clonal Cactus of the Sonoran Desert

Stenocereus eruca is a prostrated, self-incompatible cactus endemic to the Sonoran Desert that regenerates primarily through clonal propagation. Clonal growth is expected to affect mate availability by influencing the number and spatial distribution of mating types. In this paper we examine the role of clonal growth on female fecundity through a series of pollination experiments in a population of S. eruca. We set up a pollen supplementation experiment using five distance treatments with pollen collected at 1, 10, 100, 1000 and 25000 m from receptor flowers during the years 2001 and 2002 and evaluated genetic sifmilarities between pairs of receptor-donor ramets through RAPD markers. Our data on fruit set, number of seeds/fruit, germination and overall fecundity revealed that S. eruca show a significant reduction in female fecundity when pollination occurs between ramets located at short distances (1 and 10 m), while genetic data showed high levels of similarity at those distances. The reduction in female fecundity is apparently a consequence of geitonogamy and inbreeding depression. Our data suggest that clonal growth and geitonogamy are likely to be partially responsible for the low levels of sexual reproduction and seedling recruitment observed in populations of S. eruca. Co-ordinating editor: H. Kudoh     

The effects of disturbance on a herbivore-epilithon interaction in an upland stream

Two experiments were performed in an upland stream to determine the effects of the frequency of physical disturbance on the relationship between an abundant glossosomatid trichopteran (Agapetus monticolus; Banks) and the epilithon upon which it feeds. Artificial cobbles with an established epilithic community were tumbled either every 1, 2 or 4 weeks. The first experiment failed to detect any significant effects of rock tumbling on the abundance of A. monticolus or the epilithon: a result due to several spates. The first experiment did reveal that disturbances may disrupt the ability of A. monticolus to locate patches of abundant food. The second experiment found that although the abundance of A. monticolus was not affected by the disturbances, periphyton abundance was significantly reduced. Increasing the frequency of disturbance did not magnify this effect. Comparisons of these results with other studies of disturbance in streams indicate that the effects of disturbance on herbivory may be highly variable. A variety of factors, such as the relative resistances of the herbivores and the epilithon, need to be examined before the effects of disturbances on lotic herbivorous interactions can be completely understood.     

The response of hyporheic invertebrate communities to a large flood in the Hunter River, New South Wales

Previous studies on recovery in hyporheic communities have found that communities rapidly return to pre-disturbance levels. However, most of these studies have concentrated on small floods or ones with short return periods. I studied the impact of a large 1 in 6 year flood on the hyporheic community at 2 sites in the Hunter River, a large coastal river in New South Wales with a mean daily flow of 15 m³ s⁻¹. The flood peaked at 1270 m³ s⁻¹ and afterwards invertebrate densities at the 2 sites were 83 and 67% less than they were before. Recovery to pre-flood densities was slow but was aided by increases in the oligochaete and cyclopoid populations. At Site 1, there was a boom in oligochaete and cyclopoid numbers 61 d after the flood, but the communities resumed their pre-flood densities by Day 139. Recovery at Site 2 took 139 d. Most groundwater taxa (stygobites) living in the hyporheic zone did not recover from the disturbance when compared to non-stygobites. Apart from Microturbellaria and the harpacticoid Parastenocaris sp., numbers of all stygobite taxa continued to decline after the flood, becoming absent after 61 d. The poor recovery of stygobites is probably due to their adaptations for survival in the relatively stable groundwater environment. This study shows that hyporheic communities are sensitive to large bed-moving floods and supports the hypothesis that ecotonal species with a strong affinity to one ecosystem can be poor at recovering from disturbances that occur in an adjacent ecosystem.     

The metabolism of organic matter in the hyporheic zone of a mountain stream, and its spatial distribution

Community respiration in hyporheic sediments (HCR) was studied in a characteristic riffle-pool-sequence of a mountain stream. HCR activity at the riffle site strongly exceeded that at the corresponding pool site with a mean ratio of 5.3. The vertical distribution of HCR activity was homogeneous in the pool, while there was a distinct maximum in the uppermost layer in the riffle. Similarly, the spatial distribution of certain fractions of particulate organic matter (POM), and their turnover, was largely determined by stream morphology. Mean annual HCR per unit area of stream bed was estimated as 1.71 g O₂ m⁻² d⁻¹. Hence, HCR contributes significantly to total heterotrophic activity in streams, thus enhancing the relative importance of heterotrophic processes in running waters containing hyporheic zones.     

The hyporheic zone as a source of dissolved organic carbon and carbon gases to a temperate forested stream

The objective of this study was to examine chemical changes in porewaters that occur over small scales (cm) as groundwater flows through the hyporheic zone and discharges to a stream in a temperate forest of northern Wisconsin. Hyporheic-zone porewaters were sampled at discrete depths of 2, 10, 15, 61, and 183 cm at three study sites in the study basin. Chemical profiles of dissolved organic carbon (DOC), CO₂, CH₄, and pH show dramatic changes between 61 cm sediment depth and the water-sediment interface. Unless discrete samples at small depth intervals are taken, these chemical profiles are not accounted for. Similar trends were observed at the three study locations, despite each site having very different hydraulic-flow regimes. Increases in DOC concentration by an order of magnitude from 61 to 15 cm depth with a corresponding decrease in pH and rapid decreases in the molecular weight of the DOC suggest that aliphatic compounds (likely organic acids) are being generated in the hyporheic zone. Estimated efflux rates of DOC, CO₂, and CH₄ to the stream are 6.2, 0.79, 0.13 moles m² d^-1, respectively, with the vast majority of these materials produced in the hyporheic zone. Very little of these materials are accounted for by sampling stream water, suggesting rapid uptake and/or volatilization.     

Seasonal dynamics of nitrogen cycling for a Prosopis woodland in the Sonoran Desert

Prosopis woodlands in the Sonoran Desert have levels of above-ground biomass and productivity much higher than those predicted for desert plant communities with such low levels of precipitation. A stand ofP. glandulosa near the Salton Sea, California, has 13,000 kg ha^?1 aboveground biomass and a productivity of 3700 kg ha^?1 yr^?1. Such a high level of productivity is possible because Prosopis is decoupled from the normal limiting factors of water and nitrogen availability. Soil nitrogen contents for the upper 60 cm of soil beneath Prosopis canopies have 1020 g m^?2 total nitrogen, 25 per cent of which is in the form of nitrate. Such accumulations of nitrogen may be the result of active symbiotic nitrogen fixation. Early estimates suggest that about 25–30 kg N ha^?1 yr^?1 is fixed in these stands. Since Prosopis covers only 34% of the ground surface and its water resources are not limiting, much higher levels of nitrogen fixation and productivity may be possible in managed stands at greater densities.     

Evaluating Survey Methods for Monitoring a Rare Vertebrate,the Sonoran Desert Tortoise

Abstract: Effective conservation requires strategies to monitor populations efficiently, which can be especially difficult for rare or elusive species where field surveys require high effort and considerable cost. Populations of many reptiles, including Sonoran desert tortoises (Gopherus agassizii), are challenging to monitor effectively because they are cryptic, they occur at low densities, and their activity is limited both seasonally and daily. We compared efficiency and statistical power of 2 survey methods appropriate for tortoises and other rare vertebrates, line-transect distance sampling and site occupancy. In 2005 and 2006 combined, we surveyed 120 1-km transects to estimate density and 40 3-ha plots 5 times each to estimate occupancy of Sonoran desert tortoises in 2 mountain ranges in southern Arizona, USA. For both mountain ranges combined, we estimated density to be 0.30 adult tortoises/ha (95% CI = 0.17–0.43) and occupancy to be 0.72 (95% CI = 0.56–0.89). For the sampling designs we evaluated, monitoring efforts based on occupancy were 8–36% more efficient than those based on density, when contrasting only survey effort, and 17–30% more efficient when contrasting total effort (surveying, hiking to and from survey locations, and radiotracking). Occupancy had greater statistical power to detect annual declines in the proportion of area occupied than did distance sampling to detect annual declines in density. For example, we estimated that power to detect a 5% annual decline with 10 years of annual sampling was 0.92 (95% CI = 0.75–0.98) for occupancy and 0.43 (95% CI = 0.35–0.52) for distance sampling. Although all sampling methods have limitations, occupancy estimation offers a promising alternative for monitoring populations of rare vertebrates, including tortoises in the Sonoran Desert.