managed flooding for riparian ecosystem restoration
TRANSCRIPT
Managed Flooding for Riparian Ecosystem Restoration
Managed flooding reorganizes riparian forest ecosystems along the middle Rio Grande in New Mexico
Manuel C. Molles Jr., Clifford S. Crawford, Lisa M. Ellis, H. Maurice Valet!, and Clifford N. Dahm
AcenturieS-Old Indian pueblo named Isleta stands on a small rise of land west of the Rio
Grande in central New Mexico. Its name, which means "island" in Spanish, reflects the fact that the pueblo was on an island in the middle of the Rio Grande when it was first seen by Spanish explorers (Pearce 1965), The name Isleta comes from the time before flood control on the Rio Grande, when spring flood waters roared down from the San Juan Mountains of Southern Colorado, flooding the middle Rio Grande Valley of central New Mexico (Figure 1). During some years, high spring runoff would extensively flood the middle Rio Grande Valley, sending most valley residents fleeing to surrounding high ground until the flood waters receded. However, these floods rarely touched Isleta Pueblo, only transforming it into the island its name suggests. While the residents of other settlements huddled in tent villages on the surrounding mesas, the life of Isleta's residents went on as usual. Today, dams and levees built on the Rio ------~----
Manuel C. MoUes Jr. (molles@sevilleta. unm.edu) is a professor, Clifford S. Crawford ([email protected]) is a professor emeritus, Lisa M. Ellis (lellis@ sevilleta.unm.edu) is a graduate student, and Clifford N. Dahm (cdahm@sevilleta. unm.edu) is a professor in the Department of Biology, University of New Mexico, Albuquerque, NM 87131. H. Maurice Valett ([email protected]}isanassistantprofessor in the Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061. © 1998 American Institute of Biological Sciences.
September 1998
Out of this search
for general ecological
principles will come
a better understanding
of the structure
and function of
riparian ecosystems
Grande have reduced the frequency and intensity of flooding; as a result, Isleta Pueblo is surrounded not by flood waters each spring but by alfalfa fields and suburban development.
The flood-pulse concept IJ unk et al. 1989, Bayley 1995) emphasizes that water-land interactions create and maintain river-floodplain ecosystems as some of the most productive and diverse ecosystems in the world. However, river regulation has largely eliminated the flood pulse from most large rivers. Postel et al. (1996) estimated that humans now use a little over half of accessible river runoff worldwide. Fragmentation of river channels by dams, inter basin diversion, and irrigation strongly or moderately affect 77% of the total discharge of the 139 largest rivers in the northern third of the earth (Dynesius and Nilsson 1994). Benke (1990) noted that, except for the Yellowstone River in Montana, all large rivers in the 48 contiguous United States have been severely altered for flood control,
hydropower, or navigation. He also estimated that of the 5,200,000 km of streams and rivers in the contiguous 48 states, only approximately 2% are classified as high quality. Moreover, only 42 free-flowing river segments longer than 200 km in length remain. There are currently 75,000 dams on the streams and rivers of the United States (Meyer 1996), and large dams are being completed at an estimated rate of 500 per year worldwide (Covich 1993). Thus, the extensive disconnection of rivers from their floodplains represents global-scale ecological change, the consequences of which we do not understand.
Studies of the effects of modified river flow on riparian ecosystem structure and function require longterm flow records. The flow record for the middle Rio Grande is exceptionally long. The first stream gauge established by the US Geological Survey was constructed on the Rio Grande at Embudo, New Mexico, in 1889. Since its construction, the Embudo gauge has been measuring the hydrologic regime of the Rio Grande. The most salient feature of that regime is a consistent annual peak in flow at the end of Mayor the beginning of June (Slack et al. 1993). Although flows still peak at the end of May, dams on the Rio Grande have eliminated most flooding. Historically, floods beginning in May would sometimes continue until late July. This spring flood pulse made the middle Rio Grande Valley a mosaic of river channels, marshes, wet meadows, and riparian forests of
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Figure 1. A map of the middle Rio Grande Valley in New Mexico showing the location of the study area at Bosque Del Apache National Wildlife Refuge.
various ages. T aday, however, the middle Rio Grande no long-er meanders freely across its valley but is constrained to flow within its levees. Most riparian for-ests are now con-fined to the area be-tween the levees, and most of the wetlands in the middle Rio Grande have disap-peared (Crawford et al. 1993, 1996a).
What are the ecological consequences of eliminating the
co
annual flood pulse in the middle Rio Grande Valley? One of the most obvious consequences has been a dramatic reduction in the germination and establishment of native cottonwoods and willows (Howe and Knopf 1991). The riparian forest along the middle Rio Grande in central New Mexico is the most extensive cottonwood-willow forest left in the south-
Rio Puerco
Rio Salado
Cochiti Reservoir
Jemez River
Cochiti Reservoi
Albuquerque
Bosque del ApacheNWR
western United States (Crawford et a1. 1993). However, this extensive forest is largely an ecological legacy of past flooding. These stands may be rapidly senescing, and because of flood control, new stands of cottonwood are not being established. In addition, a stabilized river flow appears to favor invasion by non-native tree species, such as saltcedar,
Figure 2. A patch of riparian forest at Bosque Del Apache National Wildlife Refuge in ] 991. This patch, which had not flooded for over 50 years, accumulated large amounts of woody debris. The absence of a flood pulse may contribute to such accumulations.
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T amarix ramosissima, and Russian olive, Elaeagnusangustifolia (Campbell and Dick-Peddie 1964, Crawford et al. 1996a).
Students from the Department of Biology at the University of New Mexico have been monitoring several stands of riparian forest, or "bosque," as riparian forests along the Rio Grande are known, for approximately ten years. Monitoring of these riparian forests was prompted by the concern that without longterm data sets, these natural areas could not be managed properly. The riparian forest monitored by these students was established mainly during the last large flood on the middle Rio Grande, which took place in 1941-1942. Now, however, most of the forest along this reach no longer experiences overbank flooding. In the absence of flooding and with the invasion of non-native trees, these tracts of riparian forest have become tangled with woody debris (Figure 2). The organic matter that has accumulated on the floor of riparian forests along the middle Rio Grande now averages over 50 x 106 g/ha in some areas (MoUes et al. 1996), which increases fire danger (Stuever 1997) and immobilizes substantial quantities of plant nutrients.
Monitoring has produced several data sets , including leaf fall, which has been used as an index of forest production. Autumn leaf fall has been measured in three sections of a 760 m long segment of riparian forest within the Rio Grande Nature Center in Albuquerque, New Mexico. From 1989 to 1996, leaHall dropped steadily, declining by over 40% in just eight years. This record of leaf fall quantifies what had previously only been suspected: The bosque within Albuquerque is senescing rapidly (Figure 3). This finding gives a sense of urgency to research on riparian restoration along the middle Rio Grande.
Little is known about the potential importance of flooding for maintaining established riparian forest. To investigate how the absence of a flood pulse on the middle Rio Grande has affected the structure of microbial and animal populations on the forest floor and how the flood pulse affects the rates of ecosystem processes such as forest floor respira-
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tion and decomposition of plant litter, we created a series of managed floods in a riparian forest at Bosque del Apache National Wildlife Refuge near San Antonio, New Mexico. A major objective of our research is to evaluate the potential of managed floods as a tool for riparian restoration by measuring key population, community, and ecosystem responses to the reestablishment of flooding. In this article, we present some of our findings that bear on this approach to riparian restoration.
Restoration and managed flooding
Figure 4 presents a conceptual model of the present status of and restoration efforts in the Rio Grande bosque. The present riparian ecosystem is a complex mosaic of forest patches dominated by either non-native vegetation, represented in Figure 4 by saltcedar, or native cottonwoods. These patches can be further subdivided into those that are disconnected from the river and are subject to long interflood intervals, and those that remain connected to the river and are subject to short interflood intervals. Current restoration efforts invalve two main approaches. The first is aimed at converting non-native forests to native forest by uprooting saltcedar and establishing cottonwood forests. This approach is indicated by the arrows labeled "1" in Figure 4. The second approach is to use managed flooding to reconnect disconnected riparian forests to the Rio Grande-that is, to reduce interflood intervals. It is this restoration approach (indicated by the arrow labeled "2" in Figure 4) that we explore in our studies.
Two study sites were established in 1991 in mixed cottonwood forest at Bosque del Apache National Wildlife Refuge, elevation 1400 m, approximately 5 km south of San Antonio, New Mexico. The two sites, which lie outside the river levee, have been isolated from flooding by the Rio Grande for more than 50 years. One site was designated as a control site and the other as the experimental flood site. Although we flooded approximately 10 ha of forest, ecological monitoring was organized around a 3.1 ha, 200 m diameter
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Figure 3. Annual leaf fall at the Rio Grande Na-ture Center in Albuquer-que, New Mexico, from 1989 to 1996. Leaf fall declined by approxi-mately 40% over that period. The absence of a flood pulse in the mid-die Rio Grande may reduce riparian forest leaf production.
mammal-trapping web. In 1994, we established two additionalstudy sites with-
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in the levees at Bosque del Apache. One of these additional sites seldom floods and so served as a second dry control site. The other site has continued to flood frequently because of its topography and position within the levees and was therefore designated as the natural flood site.
The canopy at all four study sites was dominated by Rio Grande cottonwood, Populus deltoides ssp. wislizenii. ranging from 8 to 15 min height. The forest subcanopy consisted of Goodding willow, Salix gooddingii, and saltcedar. The dominant understory shrubs were seepwillow, Baccharis glutinosa. New Mexico olive, Forestiera neomexicana, and Russian olive.
We collected baseline data for two years and then flooded the experimental flood site for approximately one month during each of the following three years (Figure 5). These managed floods were designed to simulate the conditions of a lowenergy flood near the edge or in the backwater of a major flood and to saturate riparian soils and soak organic matter on the forest floor. Floods, which took place from midMay to mid-June of 1993, 1994, and
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1995, were timed to coincide with the historic flood peak of May 31 at the US Geological Survey's Embudo gauging station on the upper Rio Grande (Slack et al. 1993). Floodwater was diverted from a nearby irrigation canal that carried a mixture of water diverted directly from the Rio Grande, irrigation return flows, and groundwater drainage accumulated in the nearby Rio Grande Low Flow Channel. The level of water diverted onto the experimental flood site was controlled by preexisting berms and by water-control gates constructed by personnel of the Bosque del Apache National Wildlife Refuge at the inflow and outflow of the experimental flood site. Because the topography is uneven, water depth during managed floods ranged from approximately 20 cm to over 200 cm, with an average floodwater depth of approximately 50 cm.
An ecosystem reorganization model for riparian restoration
Although Figure 4 depicts the reestablishment of the flood pulse to restore native forest as a direct process, reestablishing the flood pulse
Dominant Riparian Vegetation
Saltcedar Cottonwood
connected connected
Short non-native native I
Figure 4. Riparian forest restoration in the middle Rio Gr'ande Valley. Approach 1 converts non-native forest into native cottonwood forest by uprooting saltcedar and establishing cottonwood through pole planting or wet-soils management to favor cottonwood seed germination. Approach 2 reestablishes the flood pulse, reconnecting riparian forest to
the river and consequently reducing the interflood interval.
Interflood Interval disconnected disconnected
non-native native Dz I Long
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to a riparian forest long disconnected from its river will initiate a distinctive ecological phase. Based on observations made during our research, we propose that flooding previously isolated riparian forests will initiate significant reorganization of the riparian ecosystem, a process much like that proposed by Bormann and Likens (1979) for hardwood forest ecosystems undergoing succession following clearcutting. We suggest that following reestablishment of flooding along the Rio Grande, this process of reorganization will eventually return the riparian forest to a position similar to its historic state (Crawford et al. 1996b). We propose that the ecological response to the restoration of flooding will involve three phases: the initial disconnected phase, represented in this study by the two dry control sites; a reorganization phase initiated by managed floods and represented in this study by the experimental flood site; and a steady-state phase that will approximate conditions prior to flood control, represented in this study by the natural flood site.
The structure and function of the riparian forest should differ substantially during the disconnected, reorganization, and steady-state phases. For instance, Figure 6 shows our model for changes in rates of forestfloor respiration during these phases. During the disconnected phase, forest-floor respiration will be limited by moisture availability. Indeed, as a consequence of moisture limitation in the arid climate of central New Mexico, forest-floor respiration should be orders of magnitude lower during the disconnected phase than during the other two phases. We predict that in mesic regions, forestfloor respiration should be more similar during the three phases.
The model predicts that forestfloor respiration will be highest during the initial stages of the reorganization phase, when organic matter pools are at their maximum and the pool of relatively labile organic mat~ ter is still large. As reorganization proceeds, annual flooding will con~ tinue to produce pulses of intense forest-floor respiration, but the height of the respiratory peak should be progressively damped as decomposition reduces the quantity and
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quality of organic matter on the forest floor. Eventually, when the pool of organic matter consists principally of the previous season's litterfall, the respiratory peaks will level off and the system will enter the steady-state phase.
Ecological responses to managed flooding
The first floodwaters diverted onto the experimental flood site crossed land that had not been subject to the flood pulse of the Rio Grande for over half a century. The rising waters of this managed flood induced immediate responses by forest-floor animals. The forest floor at the leading edge of the floodwaters came alive with hopping crickets and running spiders. However, our measurements revealed that some of the most dramatic responses to managed flooding involved unseen microbial populations and chemical and physical processes (Ellis et a1. 1996). A broad range of ecological responses to the managed flooding conducted in this study is summarized in Table 1.
Population and community responses to flooding. Microbial populations and activity were generally increased by flooding (Table I). Indications of increased microbial activity at the experimental flood site compared to the control site included increased abundance of soil bacteria, fungi, and cellulose decomposers. Both the root lengths colonized by mycorrhizal fungi and the mycorrhizal inoculum potential were also generally higher at the experimental flood site than at the control site. Dehydrogenase activity, which indicates soil biological activity, was more than twice as high at the experimental flood site.
Flooding also restructured the forest-floor arthropod community. During the course of the study, 120 species of forest-floor arthropods were collected at the control site and 116 species were collected at the experimental flood site. Managed flooding did not appear to affect the number of forest-floor arthropod species, but it did alter the relative abundance of species. Although flooding reduced both the species richness and the abundance of ants at the experimen-
tal flood site, at least one species of arboreal ant, Crematogaster cerasi, thrived in the presence of managed flooding. By contrast, floods reduced the abundance of ground-nesting ant species, such as Monomorium mini~ mum. Flooding also reduced the abundance of two non-native terrestrial isopods, Armadillidium vulgare and Porcellio laevis. Meanwhile, populations of the native floodplain cricket, Gryllus alogus, increased with managed flooding.
In contrast to the responses by microbial and arthropod populations, three years of flooding had no measurable effect on small-mammal populations (Ellis et a!. 1997). The dominant small mammal, the whitefooted mouse, Peromyscus leucopus, was abundant at both the dry control sites and at the experimental and natural flood sites. Flooding did not affect the abundance of P. leucopus or its reproductive condition. Traps set in cottonwood trees during flooding showed that P.leucopus remained in the forest throughout the managed floods. This adaptability to managed flooding is not surprising because P. leucopus is common in floodplain forests throughout North America (Ruffer 1961, Blem and Blem 1975). This highly arboreal mouse escapes floodwaters simply by climbing trees.
Ecosystem responses to managed flooding. Large quantities of dissolved and particulate organic matter in the flooded forest created a very high biological oxygen demand, approaching that of raw sewage (Lieurance et a!. 1994). The floodwaters entering the forest held approximately 8 mg/L of dissolved oxygen (DO) but were rapidly depleted of DO by high oxygen demand. DO concentrations at 15 em above the submerged litter layer fell to undetectable levels throughout the managed floods. By contrast, at the natural flood site, DO concentrations never fell below 4 mg/L within a few centimeters of the forest floor.
We used respiration chambers placed on the forest floor to measure forest-floor respiration during two managed floods and one natural flood. During the second managed flood, respiratory rates at the experimental flood site were 300 times
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Table 1. Several population, community, and ecosystem variables at control (C) and experimental flood (EF) sites.
1993 1994 1995 Response' C EF C EF C EF
Total soil bacteria (x 107 colonieslg soil) 431~ (±6)C 401 (±4) 371 (f6) 591 (±4) 391 (±4) 662 (±6) Total soil fungi (x 103/g soil) 673 (±8) 341 (fs) 45 2 (f3) 693 (±S) 36 1,l (±4) 823 (±9) Cellulose decomposers (x 105 colonies/g 20 1 (±3) 52 2 (fs) 482 (±4) 93] (±6) 46 2 (±S) 943 (±7)
soil) Dehydrogenase ().lg • g-1 , h-1) 4.19 1 (fO.45) 9.262 (±0.62) 3.751 (±OAO) 9.42 (fO.66) Mycorrhizal colonization (% root length) lll(±4) 26l (±2) 48 3,4 (±4) 494,5 (f7) 403.4 (f6) 56 5 (is) Mycorrhizal inoculum potential 321 (±4) 50' (±4) 35 1 (±4) 562 (±9) 33 1 (±9) 56 2 (is)
(% root infection) Ants (total in 30 traps in June and August) 348 18 192 9 95 10 Isopods (total in 30 traps in June 970 320 407 49 2,335 405
and August) Crickets (total in 30 traps in June 2 5 2 84 4 9
and August) Mice (individualslha in August) 22.021 (f3.09) 25.98 1 (±3.3s) 16.741 (±l.72) 13.661 (±lAS) Log decomposition (% mass remaining) 96.06 1 (±lA8) 91.76 1 (±l.ll) 95.781 (f1.29) 84.792d (±S.07) leaf decomposition (g dry mass remaining) 2.781 (fO.07) 2.182 (±0.03) 3.321 (±0.10) 1.782 (±0.06) 3.271 (±0.06) 1.982 (±0.02)
'Statistical comparisons for microbial and fungal data are across all sites and years within each row; comparisons for mice and log and leaf decomposition are control versus experimental flood for each year only. bDifferenrly numbered superscripts indicate significant differences (F < 0.01). <Values in parentheses are standard errors. dp = 0.061
higher than at the unflooded control site. During the third managed flood, forest-floor respiration was nearly 600 times higher at the experimental flood site than at the control site. The extremely high rates of respiration in the flooded forest appear to have involved many metabolic pathways, including sulfate reduction. During all three experimental floods, the flooded forest was filled with the smell of many sulfurous compounds. However, sulfurous smells were progressively reduced with each managed flood, which suggests ecosystem reorganization. Respiration rates measured at the natural flood site were approximately one-tenth the rates measured at the experimental flood site.
Flooding increased the rate of mass loss by leaves and logs (Molles et aJ. 1995, Ellis et a!. 1996). By the second and third managed floods, mass loss by leaves at the experimental flood site was 125% higher than at the control site. However, the rates of leaf decomposition were almost identical at the experimental flood and natural flood sites. Flooding probably increases the rate of leaf mass loss through leaching of soluble substances and by increasing bacterial and fungal activity (Webster and Benfield 1986, Biirlocher 1992).
Log decay rates were calculated using the exponential model Yt = yoe-kt, where Yo is the initial log mass,
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Yt is the log mass at time t, and k is the decay rate constant. At the experimental flood site, the average rate of log decay, k, was 0.065 yo', whereas at the control site it was 0.010)'1. These decay constants yield log half-lives of 10.6 years at the experimental flood site and 69.3 years at the control site. Although the three experimental floods did not significantly reduce the quantity of organic matter at the experimental flood site, the state of decay of woody debris was more advanced at this site than at the control site (Ellis et al. 1996, Molles et al. 1996).
Patterns of ecosystem reorganization
The population, community, and ecosystem responses we identified indicate that managed flooding initiated a process of ecosystem reorganization at the experimental flood site. Figure 7 pairs the conceptual model from Figure 6 with measurements of forest-floor respiration at the control, experimental, and natural flood sites. As predicted by the model, rates of respiration were two orders of magnitude higher during the experimental flood than on the dry forest floor. Contrary to the model, however, forest-floor respiration was 50% higher during the second experimental flood than it was during the first experimental
flood. This increase may be the result of less anoxia during the second flood, which produced higher rates of organic matter processing. Also as predicted by the model, forest-floor respiration at the natural flood site was intermediate between the rates observed at the control and experimental flood sites.
The qualitative agreement between the ecosystem reorganization model and measured rates of forestfloor respiration indicates that restoration of flooding to disconnected riparian forests initiates a distinctive reorganization phase during the transition to full restoration. However, we predict that the amount of time required for reorganization will vary a great deal from one ecosystem component to another. As a consequence, some aspects of restoration will be delayed longer than others. Although long-term study and continued flooding of study sites will be necessary to document variation in the tempo of reorganization, it is possible to make some preliminary predictions.
The results of the three managed floods indicate that the time required for restoration of ecosystem components will range from less than one year to decades. For instance, the minimal response of small-mammal populations to the three managed floods suggests that restoration of flooding will not lead to a reorganization of these populations. By con-
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trast, populations of decomposer fungi, mycorrhizal fungi, and surface-active arthropods, such as ants and crickets, all showed substantial responses to the experimental floods and are therefore subject to reorganization. Both mycorrhizal and decomposer fungi doubled their activity during the course of each flood at the experimental flood site but returned to levels comparable to those at the control site within months. Thus, reorganization of fungal populations appears to take less than a year. Surface-active arthropods showed a greater response to flooding than fungi, but the response appears to have built up over the course of the three floods and not to have reached steady state. Consequently, reorganization of surface-active arthropod populations will likely take several additional years. By contrast, leaf decomposition was virtually identical at the experimental flood and natural flood sites by the third managed flood, indicating that reorganization of this process was complete.
Reorganization of forest- floor organic matter should take much longer. The orders-of-magnitude increase in forest-floor respiration in response to experimental flooding indicates a dramatic response by the forest-floor community. However, even -with this high rate of respiration during flooding, three managed floods did not decrease the amount of forest-floor organic matter at the experimental flood site (MoUes et al. 1996). The predicted half-life of 10.6 years for logs at the experimental flood site suggests that reorganization of forest-floor litter may take a decade or more.
Implications for management, policy, and research
The widespread modification of river and riparian ecosystems creates an urgent need to better understand the ecological effects of isolating riparian ecosystems from rivers and to develop methods to restore or better manage these threatened ecosystems. A growing scientific consensus identifies river and riparian restoration as priority areas for future research. Naiman et al. (1995) prioritized six research areas in freshwater ecol-
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ogy. The first priority was to develop principles to guide restoration of aquatic ecosystems and to develop methods to assess the success of restoration, the second was to determine principles for maintaining biodiversity, and the third was to explore the effects of modified hydrologic flow patterns on ecological processes. These three priorities are addressed by the effects of our managed floods on the composition of forest-floor communities and on rates of organic matter processing and by the ecosystem reorganization model for riparian restoration that we have proposed to explain these effects.
Restoration of river and riparian ecosystems has become a topic of intense interest worldwide. The managed flood on the Colorado River in the southwestern US in the spring of 1996 focused scientific and public attention on methods for managing and restoring river and riparian ecosystems through managed flooding (Collier et al. 1997; Schmidt et al. 1998). Reestablishing flooding on a river whose flow has been regulated for decades is now seen as a potential means to restore riverine fluvial geomorphology and to improve the chances for survival of threatened and endangered species in a riverine landscape. Another major restoration project is directed at the channelized and regulated Kissimmee River in South Florida. This project, which is scheduled to begin in 1998, is designed to restore 70 km of river channel and 11,000 ha of wetlands over the next 15 years (Dahm et al. 1995, Cummins and Dahm 1995, Toth et al. 1998).
Ambitious projects such as these represent historic initiatives in ecosystem restoration; however, they are a small part of the challenges that remain to restore degraded rivers and riparian zones throughout the world. Our research has yielded some of the first quantification of riparian ecosystem response to managed flooding. The observed rates of response and the ecosystem reorganization model for riparian restoration provide estimates of the potential trajectory and time required for riparian restoration with managed flooding.
Nowhere are anthropogenic impacts on river ecosystems greater than
in the arid and semi-arid regions of the world. Water is the lifeblood of these regions, which constitute approximately 33% of the land surface worldwide. Increasing human demand for fresh water in arid regions leads to inevitable conflicts between the water needs of human populations and those of native ecosystems. The mismatch between water availability and water demand has placed most rivers and riparian zones of arid lands in peril. Based on historic habitat losses and increasing demands for water in the region, Christensen et al. (1996) describe all riparian forests in the United States as threatened ecosystems and the riparian forests of New Mexico, Arizona, and California as endangered.
Research to determine the basic principles for protecting and restoring these endangered ecosystems is, therefore, urgently needed. Our own studies have examined a broad spectrum of responses to managed flooding by a riparian ecosystem ina semiarid region. Some of the results of this research have already been incorporated into policy and management recommendations for the Rio Grande (Crawford et a1. 1993, 1996b). For instance, water managers along the Rio Grande are now considering a program of managed flooding, especially during years of higher water availability, to safeguard the health of the riparian forest. With this same goal in mind, academic scientists and water managers are conducting cooperative studies to quantify the potential water costs of managed flooding of riparian forests along the middle Rio Grande in New Mexico. The results of this work along the Rio Grande may have management implications for large, regulated rivers in other semi-arid regions.
There has been a growing understanding of the functional role of riparian zones (e.g. Gregory et al. 1991, Stanford and Ward 1993, Bayley 1995), which are now recognized as areas of high primary production, exceptional biodiversity, and rapid biogeochemical cycling. Our research applies an ecosystem perspective to riparian zones, similar to that emphasized by Gregory et al. (1991) and Beneala (1993), in whieh hydrologic exchange integrates river-
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Figure 5. The experimental flood site at Bosque Del Apache National Wildlife Rcf~ uge in flood in 1994. Managed floods on this site flooded approximately 10 ha of riparian forest that had been disconnected from the Rio Grande for over 50 years.
ine and catchment systems and strongly influences biogeochemical processes and populations in riparian zones. Because flood control has limited the connections between rivers and riparian ecosystems, we are particularly interested in determining how variation in the interflood interval affects ecosystem structure and function. In most cases, reestablishing the full spectrum of historical river-riparian interactions is not a feasible management opt ion (Crawford et al. 1996b). [t appears, how· ever, that managed floods can be used ro partially restore riparian ecosystems.
It is essential to combine studies of ecosystem functional responses with community and population studies. By studying a broad spectrum of ecological responses to managed floods, we hope to resolve empirically [he time course of ecosystem reorganization in response to hydrologic manipulation. Moreover, organizing experimental studies of restoration around conceptual models is likely to lead to general principles for ecosystem restoration. For instance, our proposed model of ecosystem reorganization led to tWO insights that should be considered during riparian restoration . Firsr, during restoration, the ecosystem cannot proceed directly from the disconnected phase to the steady-state phase but must pass through a period of reorganization. Second, ecosystem components reorganize at different rates, and re stora tion projects must recognize and consider this va riation in the tempo of reorganization. Out of this search for general ecological principles will come a better understanding of the structure and function of riparian ecosystems and more effecti ve programs for their restoration.
Acknowledgments
Many people have contributed ro the studies described in this article. We thank Phil Norton, John Taylor,
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Mauged floods Ixgin ... 10,000 -r------'r------
Figure 6. The reorganization model for riparian restoration. Resto-
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Rpmli"'" (mg C nil d·1 )
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ration involves three phases: the disconnected phase, prior to the reestablishment of managed floods; the reo o rganization phase, induced by (he reestablishment of the flood pulse; and the steady-state phase, which follows the reo rganization phase. The predicted response
to restoration plotted here is forest -floor respiration (R). Dotted line includes proposed within-year variation; solid line presents predicted average annual rates.
and th e staff at Bosque del Apache a) National Wildlife Refuge for their generous logistical support. We are indebted to Shivcharn Dhillion, Tom Kieft, and Carl White for help and advice on soils and soi l ecol-ogy and to numer-
Figure 7. Predicted and observed rates of forest ~fl oor respiration. (a) Conceptual model (from Figure 6). (b ) Observed rales of forest"f1oor respira tion at concrol sites, ar the exper im en tal flood site, and at the natural flood site.
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ous undergraduate and graduate students atthe University of New Mexico. Insightful discussions with Peter Jacobson were much appreciated. This research was partially supported by Cooperative Agreement 14-16-0002-91-228 between The University of New Mexico and the US Fish and Wildlife Service, by NSF grant no. DEB- 9414767, and by NASA Ecosystern Restoration Award NAG5-6999.
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