Amencan F~sheriesSociety Symposium 37:179-193, 2003

Fish Relationships with Large Wood in Small Streams

USDA Forest Service, Southern Research Station, Department ofFisheries and Wildlife Virginia Tech, Blacksburg, Virginia 24060, USA

USDA Forest Service, Southern Research Station 1000 Front Street, Oxford, Massachusetts 38655, USA

Abstracf.-Many ecological processes are associated with large wood in streams, such as forming habitat critical for fish and a host of other organisms. Wood loading in streams varies with age and of riparian vegetation, stream size, time since last disturbance, and history of land use. Changes in the landscape resulting from homesteading, agriculture, and logging have altered forest environments, which, in turn, changed the physical and biological characteristics of many streams worldwide. Wood is also important in creating refugia for fish and other aquatic species. Removing wood from streams typically results in loss of pool habitat and overall complexity as well as fewer and smaller individuals of both coldwater and warmwater fish species. The life histories of more than 85 species of fish have some association with large wood for cover, spawning (egg attachment, nest materials), and feeding. Many other aquatic organisms, such as crayfish, certain species of freshwater mus- sels, and turtles, also depend on large wood during at least part of their life cycles.

Introduction Because decay rate and probability of displace- ment are a function of size, large pieces have a Large wood can profoundly influence the struc- greater influence on habitat and physical processes ture and function of aquatic habitats from head- than small pieces. In general, rootwads, branches, waters to estuaries. In streams and flowing snags, detached boles, and other pieces 1.0-1.5 m through forests worldwide, large wood slows, long and wider than 10 cm in diameter are de- stores, and redirects surface water and sediments fined as large wood. Although somewhat arbi- and provides cover, substrate, and food used by trary, this size approximates the wood that is en- fish and other vertebrates, aquatic invertebrates, trained rather than simply p;$sed through and microbes (Table 1).We here review the role of channels of small perennial streams. Smaller large wood in small streams and highlight spe- pieces and fragments, even those too small to cific examples of how wood influences the distri- count, also play a role in forming habitat (for ex- bution, abundance, and life history of various or- ample, as part of the matrix of material that forms ganisms. Our focus in this chapter will be on small debris dams; Bilby and Likens 1980; Dolloff and streams, defined here as fourth-order or lower and Webster 2000). Depending on the characteristics having a bank-full width of 10 m or less. of the stream channel (for example, the presence of hydraulic controls, channel constrictions, chan- nel roughness) and relative size and quality (for Physical Effects in Stream example, surface area and resistance to decay) of Channels the wood, wood accumulations or dams of both large and small wood create habitat diversity and Woody material of all sizes-from tiny fragments may persist for a few months to hundreds of years to intact trees-plays a role in stream systems. (Bilby and Likens 1980).

180 DOLLOFF AND WARREN

TA~LE1. The physical and biological roles of large wood in small streams. Physical Biological Water Substrate Food Cover storage storage substrate protect from predators quality routing abundance isolate competitors availability protect from displacement Channel shape delivery width, depth substrate habitat type type

Nearly all wood that impinges on stream multiple times, in the last 150 years. Most have channels has the capacity to influence habitat and not had time to produce the large trees that are aquatic communities. Large wood oriented per- not only the source of instream wood, but are also pendicular to the thalweg is often associated di- critical in forming complex, long-lasting habitat rectly with pool formation (Cherry and Beschta configurations (Dolloff and Webster 2000). The 1989; Richmond and Fausch 1995; Hauer et al. loss of old or late-successional riparian forests in 1999). Even wood outside or above the wetted the last century constitutes a new disturbance re- portion of the channel influences pool formation gime. The drastically reduced recruitment of large by directing patterns of scour at bank-full flows. wood coupled with natural processes of decay and Wood bridges eventually weaken with decay, downstream transport have resulted in unnatu- break into two or more pieces, and exert entirely rally low accumulations of large wood in streams different influences on channel shape and habitat across entire landscapes. Recovery of stream eco- formation. systems through formation of wood dams may The amount of large wood in streams (wood not occur for many decades (Golladay et al. 1989; . loading) varies with age of the riparian forest, Webster et al. 1990). The absence of large wood in species of riparian vegetation, size of stream, time riparian forests over much of North America has since disturbance, and land-use history. All of greatly reduced habitat complexity in streams and these factors interact with the frequency and mag- also reduced variability among streams draining nitude of natural disturbances as well as human a given ecoregion or geoclimactic setting. With- perturbations. Of these human alterations, history out adequate large wood, these systems become of land use, particularly urbanization, agriculture, more homogeneous with reduced habitat com- and logging, has changed not only the physical plexity, species diversity, and productivity. and biological characteristics of streams but also Habitats in small streams are particularly sen- our perception of what is natural. Natural distur- sitive to changes in large wood loading, and the bances like fire, wind, and floods do not typically full implications of reduced amounts of wood over affect all parts of a landscape equally. A flood- large areas are unknown but potentially far- producing storm in one watershed may not simi- reaching for certain groups and life stages of or- larly affect an adjacent watershed. Because of the ganisms. Risk to the persistence, distribution, and dynamic spatial effects of natural disturbance re- abundance of pool-dependent species may be gimes, large wood loading and hence stream habi- compounded after a natural disturbance, particu- tat features across natural landscapes vary greatly. larly if the abundance of suitable alternative habi- At any one time, some stream corridors may have tats is already low or fragmented because of a large amounts of large wood configured into previous disturbance. highly complex habitats, but others-perhaps Directly removing wood typically results in even in the same watershed-may lack wood and loss of pools and habitat complexity; in sand-bed have greatly simplified habitats (Sedell et al. 1990). streams, it may trigger or exacerbate destructive Today, the lack of large wood in streams near channel incision (Shields and Smith 1992).Remov- urban centers or in those draining agricultural ing wood from the channel or reduces regions is readily apparent, but even streams flow- numbers, average size, and biomass for both ing through forested regions often lack significant warmwater (Angermeier and Karr 1984; Shields amounts of wood (Maser and Sedell1994).Nearly et al. 1994) and coldwater (Elliott 1986; Dolloff all of the riparian forests across the eastern United 1986) fish species. Adding wood typically in- States have been harvested at least once, if not creases total pool area and depth, but simply add- FISH REIATIONSFIIPS WITH LARGE WOOD IN SMALL STRFAMS 181

ing wood may not produce the same result in all termediate-size streams (second to fourth-order) streams. The quality of habitat produced by large tend to be larger, more variable in size, and wood is a function of geomorphic characteristics, clumped at or near bends or changes in channel such as channel size, channel form, gradient, and gradient. The mechanism of entry also influences substrate size. For example, pool habitat more the arrangement of large wood accumulations. In than doubled within 1 year of adding large wood contrast to the relatively loose architecture of to a small, low gradient (3.0% slope) Virginia wood jams that form incrementally from the trans- stream (Hilderbrand et al. 1997). In contrast, pool port and entrainment of individual pieces in a area remained essentially unchanged after simi- stream, large wood pieces from landslides, debris lar amounts of large wood were added to a mod- flows, and snow avalanches tend to deposit in erate gradient (3-6%) stream where boulders were interlocking wood accumulation's (see Benda et the dominant pool-forming element. al. 2003, this volume). Although formed by dif- Wood is deposited and accumulates in small ferent processes, both types of accumulations streams by a combination of chronic and episodic make important contributions to habitat. processes (Table 2). Inputs of the greatest amounts Some large wood accumulations in small of large wood usually can be traced to specific, streams last for hundreds of years. However, the weather-related events; piles of logs deposited by total amount of large wood is dynamic, chang- floodwaters along stream sides are obvious indi- ing, sometimes dramatically, over the course of a cators of the importance of episodic events. But year. As an illustration of this variability, the half- for perspective, the annual input of organic ma- life of individual pieces of large wood ranges from terial to small upland streams from seasonal lit- 2.3 to 230 years (Harmon et al. 1986). The half-life ter fall (leaves, twigs, small branches) typically of wood dams in very small streams tends to be xceeds the input of large wood (Webster et al. less, about 6-24 months, depending on size and 990).In general, large wood input rates vary with composition of the material in the dam and char- ee species and age, slope and shape of surround- acteristics of flow. Whether limbs or whole trees, g land forms, health and history of surround- decay begins with death. The longevity of intact g forest (insects, disease, land use), and size of pieces of large wood is thus influenced by decay e watershed. Recruitment rates are generally rates (resistance to abrasion, breakage, and rot) ower in second-growth forests; in eastern North and transport processes. Softwoods generally de- America, input rates tend to be very low because cay faster than hardwoods, but certain softwood many riparian forests have been completely re- species, like hemlock, that contain high levels of moved or have been harvested repeatedly (three decay-resistant tannins and other substances, last or more timber rotations). longer than many "soft" hardwoods, such as tu- Hydraulic processes control the particular lip poplar and aspen. Temperature of both the air arrangement of large wood after it enters a stream and water, exposure to sunlight and humidity, and channel, but the mechanism of entry can also be frequency and duration of flow all affect the rate influential. Stream size, which incorporates dif- of wood decay and its longevity at a particular ferences in flow magnitudes, has a major influ- site. Wood buried in the streambed or continu- ence on the fate and distribution of large wood ally submerged resists decay better than wood (Bisson et al. 1987). In small streams (first- and that is only partly or periodically submerged, al- second-order), the dominant spatial pattern ap- ternating between being wet and dry. pears random: dams may form at predictable in- The loading of large wood in streams flow- tervals from leaves and litter; however, large ing through forests varies greatly, from 2.5 to 4,500 pieces tend to stay where they fall because trans- m3/ha (Harmon et al. 1986). In general, streams port power is low. Accumulations of wood in in- flowing through forests undisturbed by human

TABLE2. Mechanisms of large wood input to small streams. Chronic processes Episodic processes Litterfall, self-pruning from riparian areas Tree fall from stream bank failures Mortality of trees from insects and disease Tree fall from windthrow Tree fall from streambank undercutting Snow and ice accumulation and melting Avalanches, debris flows 182 DOLLOFF AND WARREN

activities have the greatest volumes or density of and larger fish than shallower areas because of large wood. High loads are well documented in the greater space (volume) of habitat available, the Pacific Northwest, but small streams flowing particularly during times of exceptionally low through forests in eastern North America also flow, such as in late summer or winter. Streams have substantial loads (Hedman et al. 1996). By with poorly developed, shallow pools have a low inference, historical amounts of large wood in residual pool volume (Lisle and Hilton 1992) and many small eastern streams probably were high. consequently tend to support fewer species, have In the southern Appalachians, streams draining simple trophic structure, and show higher vari- old-growth forests contained amounts of large ability in fish abundance (Schlosser 1987). wood comparable to those in northwestern old- Pools can form around any obstruction that growth forests and up to four times more large creates friction and resists displacement by flow- wood than in eastern wilderness streams drain- ing water, and large wood is often the dominant ing forests cleared early in the century (Flebbe and pool-forming element. In small streams in forested Dolloff 1995). Likewise, small streams in unlogged areas, large wood can be responsible for 50-9076 of watersheds of the Great Smoky Mountains Na- all pools (Andrus et al. 1988; Hedman et al. 1996). tional Park contained 4 times the volume of wood The proportion of pools formed by large wood and 10 times the material in wood dams than tends to be much lower (<1O0/0) in streams flowing streams logged early in the 20th century (Silsbee through second-growth and younger forests. and Larson 1983; Table 6 in Harmon et al. 1986). Large wood is an important habitat that is In small streams of southern New Zealand, re- included in the definitions of many fluvial habi- searchers determined that the amount of large tat types (Bisson et al. 1982; Hawkins et al. 1993; wood in older forest streams was greater than in ~rmintrout1998). Large wood can contribute to young forests, with overall amounts much smaller pool formation in a variety of ways depending relative to those found in North American forested on factors such as the size of the piece, the mecha- streams (Evans et al. 1993). nism of entry into a stream channel, channel size and gradient, and streamflow. The most corn- monly envisioned orientation is channel spanning Habitat Relations and perpendicular to the channel, which often results in forming plunge pools in the down- For many aquatic organisms, particularly fishes, stream and dam pools in the upstream direction. large wood is most important in creating and Perpendicularly oriented large wood that juts into maintaining deepwater or pool habitat. Pools and but does not span a channel may create eddies other habitats associated with large wood are at- and backwater pools. The deepest pools form be- tractive to fish for a variety of reasons. Conspicu- hind pieces that span the entire width of the chan- ous among them are the lower water velocities nel and are oriented perpendicular to flow. Large and greater depths associated with pools during wood helps maintain the diversity of physical low-flow periods. Salmonids and drift-feeding habitat by delimiting pool position and increas- minnows inhabit areas with low velocity water ing depth variability. and make forays into fast water to capture food Although the role of large wood in deep pool items in the drift (Matthews 1998).Trout and min- formation is important, its contribution as an ele- nows of eastern England were also found to be ment of channel roughness in shallow stream strongly associated with large wood (Punchard reaches should not be overlooked. In many coastal et al. 2000). Several American species also seek plain streams of the eastern United States, riffles the lower velocities behind logs to avoid swift, and cobble substrate are absent and gravel is rare cold winter flows. Refuges from high velocities, (Meffe and Sheldon 1988). Here, large wood is such as those found in the lee of logs, may sig- often the only element contributing to channel nificantly affect overwintering survival of the fed- roughness and, hence, to the forming of complex, erally listed Efheostoma rubrum (Ross flowing habitats (Smock and Gilinsky 1992). The et al. 1992) and many other small benthic fishes, presence of wood accumulations, even relatively particularly in sand-bottomed coastal plain small-diameter pieces, in shallow, sandy, flowing streams. Still other fishes use the cover of logs in areas creates a heterogeneous zone of variable pools for nest sites and spawning (for example, velocities and depths. These wood-formed "riffles centrar-chids) or attach their eggs to logs (for ex- and runs" in turn support a significant propor- ample, shiners Cyprinella spp. and relict darter tion of the stream fish diversity in coastal plain chienense). Pools typically harbor more FISH RELATIONSHIPS WITH LARGE WOOD IN SMALL STREAMS 183

streams and likely are critical to the persistence physical habitat partitioning. Angermeier and of many darters ( spp. and ~theo>tomaspp.) Karr (1984) experimentally removed large wood and Noturus spp. (Chan and from one side of a reach of a small Illinois stream Parsons 2000; Warren et al. 2002). and found that, in addition to more and larger Large wood creates sediment storage sites fish, the side of the stream with wood had more ( and terraces), the wetted portions of litter and benthos. In coastal plain headwater which contribute to producing food (algae and streams of Virginia, invertebrate densities were macroinvertebrate) and spawning habitat. Form- 10 times and biomass 5 times higher in wood dams ing new sediment terraces is particularly impor- than on sandy substrates (Smock et al. 1989). In tant in drainages where natural floodplain depos- pumice-bed streams of New Zealand, summer its and terraces have been lost, such as in many densities of macroinvertebrates were higher in small watersheds in the Cascade Range of Oregon streams where wood was present, providing ei- and across much of eastern North America. Con- ther increased habitat or greater food resources versely, large wood also encourages scour and or a combination of both (Collier and Halliday promotes transportation of fine sediments, 2000). Wood snags in southeastern Australian thereby exposing gravel where fish spawn and lowland streams serve as important habitat for benthic macroinvertebrates thrive. In general, macroinvertebrates, influencing not only the di- streamflow determines the influence of large versity of species, but also their abundance wood on routing and rates of sediment scour and (O'Connor 1991). Invertebrates associated with fill. At low flow, pools formed by wood tend to wood substrates form essential elements of the fill and riffles tend to scour, but when flow is high, diet of many fishes in these streams (Felley and pools scour and riffles are depositional. Felley 1987). Although many researchers have focused on Aside from its role in creating pools and ve- how large wood affects physical processes in locity breaks, large wood can be in important streams, large wood also has a major impact on source of cover and habitat complexity for aquatic many, if not all, biological processes, such as sub- species. Wood cover can be simply an overhang- strate type, water depth, and velocity. These physi- ing log or intricute accumulation of wood and cal processes are influenced by large wood. Large other material, often referred to a wood jam. In wood also facilitates primary production by pro- northern Japanese streams, masu salmon Onco- viding attachment sites for microbes and algae, rkynchus masou density was correlated with wood sources of nutrients, and storage zones for organic cover in both pool and nonpool habitats (Inoue matter. Biofilms, composed of microorganisms and Nakano 1998). In general, streams with com- and the mucoidal matrix that surrounds them, plex habitats tend to have more fish and fish spe- develop readily on wood, a substrate that also cies than streams lacking complexity. Complex- provides a portion of the nutrients required to ity provides a measure of security from predators maintain them (Aumen et al. 1990; Golladay and (Harvey and Stewart 1991), isolation from com- Sinsabaugh 1991; Tank and Winterbourn 1995; petitors, and points of refuge from severe envi- Benke and Wallace 2003, this volume). Wood also ronmental conditions, such as flood or drought. creates conditions favorable to the entrainment Wood provides shadow, which not only makes and processing of terrestrially derived fine par- fish harder for predators to see, but also aids fish ticulate organic material (FPOM) from riparian in seeing approaching predators (Helfman 1981). vegetation.-~icrobesand macroinvertebrates pro- Large wood protects fishes from aquatic preda- cess this material more readily than wood. Large tors (Everett and Ruiz 1993), and complexity wood also traps and retains carcasses of spent fish thwarts diving and wading predators (Power in streams that are home to runs of anadromous 1987; Harvey and Stewart 1991. Complexity is fish (Cederholm and Peterson 1985). Nutrients, critical for many fishes, particularly aggressive particularly nitrogen, from decomposing car- species like salmonids, which do not tolerate con- casses boost primary production by enhancing specifics in close proximity. Submerged branches both algal and biofilm development (Richey et al. and other wood partition habitat and visually iso- 1975; Wold and Hershey 1999). late individual fish, allowing more fish per unit Large wood enhances secondary production of available space (Dolloff 1986). Field observa- both directly, such as by increasing the surface tion of this "condominium effect" provides part area available to macroinvertebrate grazers and of the rationale for enhancing habitat by using scrapers (Benke et al. 1985), and indirectly, by discarded Christmas trees, brush bundles, and 184 DOLLOFF AND WARREN

tops from softwood trees (Boussu 1954; Reeves et regate by species and size. Particularly among fish al. 1991). with aggressive natures, physical habitat parti- Complex cover and habitat, like that created tioning is an important element of coexistence. by large wood, is especially important during Larger or more aggressive individuals tend to times of stress, such as when streamflows are ex- occupy the most efficient positjons in small ceptionally low or high (Swales et al. 1986; streams (Fausch and White 1981), which often are Pearsons et al. 1992). In small, drought-prone characterized by structure at the heads of pools, streams, large wood and other channel obstruc- near but out of high-velocity currents. Juvenile tions may provide the only refuge during low salmonids of different species and sizes are known flows. White pines planted near Benson's Run, to segregate in small streams where large wood Virginia toppled into the stream by the laterally is the dominant pool-forming element. For ex- migrating channel caused local scour and in- ample, Dolloff and Reeves (1990) observed that creased numbers and distribution of pools that coho salmon 0.kisutch and Dolly Varden S. rnalrna are used as low-flow refugia by a host of fish spe- occupied distinct regions in small (2-10-m2)pools cies, inchding brook trout Salvelinus fonfinalis in Alaska streams. One-year-old and older coho (Figure 1). Species other than fish also frequent salmon typically occupied positions at the heads these pools; predators such as raccoons, herons, of pools near the water surface, but similar-sized water snakes, and anglers are known to exploit Dolly Varden were associated more closely with the dense congregations of fish whose only es- the stream bottom and dense (frequently wood) cape lies in the tangled roots of the toppled trees. cover. Young-of-the-year coho salmon and DolIy Adult cutthroat trout 0. clarki in northern Cali- Varden occupied positions similar to the larger fornia small streams occupied habitat influenced fish but farther down stream in the pools and as- by large wood during winter floods and showed sociated less closely with cover. a strong fidelity to wood-formed pools during Multivariate studies of warmwater, stream normal fall and winter flows (Harvey et al. 1999). fish communities often demonstrate habitat par- In individual habitat units, fish tend to seg- titioning based on factors involving amount of

FIGURE1. Refuge habitat created by the rootwads of toppled strearnside trees in Benson's Run, Virginia. FISFI RELATIONSHIPS WITH LARGE WOOD IN SMALL STREAMS 185

wood and cover after depth and velocity are taken sayanus, are strongly associated with complex into account (Baker and Ross 1981; Felley and Hill wood habitats in small coastal plain streams 1983; Felley and Felley 1987; Taylor et al. 1993; (Monzyk et al. 1997; Chan and Parsons 2000) of Warren et al. 2002). The actual association of spe- the southeastern United States. Large wood is a cies-rich, warmwater fish assemblages with wood requisite feature of the diurnal cover of these spe- and cover is often a complex mix of velocity pref- cies, but specific characteristics (for example, cav- erences, behavioral responses (such as predator ity space, entrained leaves, adjacent flow) also avoidance), or feeding responses, which can only influence fish use (Monzyk et al. 1997). Brush be partially resolved in fish assemblage-habitat bundles, leaf packs, and faux rootwads experi- studies (Meffe and Sheldon 1988). In small north- mentally placed in northern streams ern Mississippi streams, many of which are af- were used by 32 species of small-stream fishes. fected by stream incision, the relative abundance Strong position effects on fish numbers and iden- of sunfishes was positively associated with wood tity reflected depth and velocity at individual sam- in relatively deep habitats, a response attenuated plers. Brush bundles and leafpacks held equal by stream incision (Warren et al. 2002). In flow- numbers of fish; smaller root samplers held fewer ing habitats, large wood was associated positively fish, but darters used them regularly. In January with the relative abundance of darters (genera (water 2-5"C), up to 70 lethargic cyprinids occu- Etheosfoma and Percina) and catfishes (mostly pied a single bundle, suggesting that such refuges Noturus spp. and bullheads Ameiurus are critical winter habitat (A. J. Sheldon, Univer- spp; Warren et al. 2002). In Ozark streams, stream- sity of Montana, M. L. Warren, Jr. and W. R. Haag, dwelling rock bass and smallmouth bass showed USDA Forest Service, unpublished data). Even in high association with wood, but habitat use the arid Southwest, large wood forms important showed distinct differences in segregation by spe- habitat for native fishes. Presence of pools with cies, size, and relation to current velocities (Probst root masses of standing or uprooted trees was the et al. 1984). Rock bass Ambloplites rupestris were best predictor of the occurrence and abundance more often associated with rootwads, regardless of the federally threatened Gila chub Gila of size, but larger smallmouth bass Micropterus infermedia (Probst and Stefferud 1994). dolomieu (>350 mm) used log complexes in higher A cursory compilation of southeastern U.S. velocities than velocities associated with rock bass fishes that have some association with large wood cover (Probst et al. 1984). is instructive (Table 3.). For many of these spe- As fish mature, their use of large wood in cies, presence of large wood is facultative, par- streams can shift in relation to their size and food ticularly where rocky substrates or other elements preferences. For example, in a prairie stream in can substitute for cover. For other species, large Kansas, growth increments of some stream fishes wood is the primary element of habitat complex- were associated strongly with amounts of large ity, especially in streams of predominantly fine wood (Quist and Guy 2001). Creek chubs Semotilus substrates. Although not exhaustive, our compi- afromaculatus were strongly associated with lation of fish associated with large wood habitat amounts of large wood for only the first year of includes 12 families and more than 86 species of growth, but all growth stages of green sunfish warmwater fishes. For these species, association Lepomis cyanellus were associated with wood (Quist with large wood includes diurnal cover, cover for and Guy 2001). The authors atkibuted the response adult or juvenile stages, food sources (such as al- of creek chubs to shifts from eating invertebrates gae or macroinvertebrates on wood), and repro- while small to eating fish when they were larger. ductive uses (such as spawning and nesting cover, Green sunfish likely exploited invertebrates on egg attachment). These species, and likely others wood at small sizes but, at large sizes, continued that live in small streams, find protection from to use large wood as cover (Quist and Guy 2001). predators, as well as resting and foraging areas Many other fish species are also associated and spawning sites among the branches and with accumulations of large wood. In Kentucky's pieces that comprise the matrix of wood jams. Beaver Creek, for example, divers found the high- Much information has accumulated showing est concentrations of the federally threatened the relation of large wood to fish populations, par- blackside dace Phoxinus cumberlandensis in pools ticularly of salmonids in western North America. with a complex mixture of large and small wood. Clear demonstration of similar relations in other Two nocturnally active fishes, the brown macltom regions and for other species, however, has often Noturus phaeus and the pirate Aphredoderzis been problematic. For example, though Flebbe and 186 DOLLOFF AND WARREN

TABLE3. Warmwater fish species associated with large wood in the southeastern United States - Species Common name Association with large wood Reference Ichthyomyzon castaneus Chestnut lamprey Spawning cover Becker 1983 Lampetra aepyptera Least brook lamprey Cover for transforming larvae Walsh and Burr 1981 Amia calva Bowfin Cover, nesting materials, Scott and Crossman 1973 nesting cover Esox niger Chain pickerel Cover Scott and Crossman 1973 CyprineIla analostana Satinfin shiner Egg attachment Gale and Buynak 1978 Campostoma spp. Stonerollers Feeding on algae Matthews 1998 Cyprinella callitaenia Bluestripe shiner Probable egg attachment Wallace and Ramsey 1981 Cyprinella galactura Whitetail shiner Egg attachment Pflieger 1997 Cyprinella lutrensis Red shiner Feeding, egg attachment Quist and Guy 2001; Pflieger 1997 Cyprinella spiloptera Spotfin shiner Egg attachment Pflieger 1965 Cyprinella venusta Blacktail shiner Egg attachment Pflieger 1997 Cyprinella whipplei Steelcolor shiner Egg attachment Pflieger 1965 Exoglossum laurae Tonguetied minnow Nesting cover Trautman 1981; Jenkins and Burkhead 1994 lossurn urn maxillingua Cutlips minnow Nesting cover Jenkins and Burkhead 1994 Opsopoeodus emiliae Pugnose minnow Probable egg attachment Johnston and Page 1990 Nocornis leptocephalus Bluehead chub Nesting cover Jenkins and Burkhead 1994 phoxinus cumberlandensis Blackside dace Cover Starnes and Stames 1978 phoxinus tennesseensis Tennessee dace Cover Starnes and Jenkins 1988 Pimephales notatus Bluntnose minnow Nesting cover, egg attachment Hubbs and Cooper 1936; Scott and Crossman 1973 Pimephales promelas Fathead minnnow Nesting cover, egg attachment Wynne-Edwards 1932; Scott and Crossman 1973 Pimephales vigilax Bullhead minnow Nesting cover, egg attachment Parker 1964 pteronotropis euryzonus Broadstripe shiner Cover Mettee et al. 1996 semotilus atromaculatus Creek chub Feeding (juvenile), diurnal Pflieger 1997; Quist and cover Guy 2001 Erimyzon oblongus Creek chubsucker Cover Mettee et al. 1996 Erimyzon tenuis Sharpfin chubsucker Cover Mettee et al. 1996 Ameiurus melas ' Black bullhead Nesting cover Breder and Rosen 1966 Ameiurus natalis Yellow bullhead Nesting cover Carlander 1969 Ameiurus nebulosus Brown bullhead Nesting cover Blumer 1985 Ameiurus serracanthus Spotted bullhead Cover Mettee et al. 1996 Noturus flavater Checkered madtom Cover Pflieger 1997 Noturus flavipinnis Yellowfin madtom Diurnal cover Dinkins and Shute 1996 Noturus finebris Black madtom Cover Mettee et al. 1996 Noturus gyrinus Tadpole madtom Cover, nesting cover Burr and Warren 1986; Burr and Stoeckel 1999 Noturus gilberti Orangefin madtom Nesting cover Burr and Stoeckel1999 Least madtom Cover, nesting cover Mayden and Walsh 1984; Etnier and Stames 1993 Noturus leptacanthus Speckled madtom Cover, probably nesting cover Mettee et al. 1996 Noturus miurus Brindled madtom Cover, nesting cover Burr and Warren 1986; Mettee et al. 1996; Burr and Stoeckel 1999 Noturus mzinitus Frecklebelly madtom Cover Etnier and Starnes 1993 Noturus nocturnus Freckled madtom Cover Burr and Warren 1986; Mettee et al. 1996 FIS~IRELATIONSHIPS WITH URGE WOOD IN SMALL STREAMS 187

TABIE3. Continued Species Common name Association with large wood Reference Noturus phaeus Brown madtom Diurnal cover, probably Monzyk et al. 1997 nesting cover Nofurus stigmosus Northern madtom Cover Etnier and Starnes 1993 Pylodictis olivaris Flathead Cover, nesting cover Jackson 1999 Aphredoderus sayanus Pirate perch Cover, feeding Benke et al. 1985; Monzyk et al. 1997 Fundulus olivaceus Blackspotted Cover Mettee et aI. 1996 topminnow Lucania goodei Bluefin killifish Cover Mettee et al. 1996 Cotttls bairdi Mottled sculpin Nesting cover, egg attachment Rohde and Arndt 1982 Arrrbloplites ariommus Shadow bass Cover Mettee et al. 1996 Ambloplites rupestris Rock bass Cover Probst et al. 1984; Matthews 1998 Centrarchus macropterzls Flier Cover Mettee et al. 1996 Enneacanthus gloriosus Bluespotted sunfish Cover Mettee et al. 1996 Leponlis spp. Cover (various life stages), Benke et al. 1985; feeding Matthews 1998 Micropterus dolomieu Smallmouth bass Cover Probst et al. 1984 Etheostoma asprigene Mud darter Cover, probable egg Page et al. 1982; Burr and attachment Warren 1986; Pflieger 1997 Etheostoma artesiae Redfin darter Cover Mettee et al. 1996 Etheostoma boschungi Slackwater darter Cover Etnier and Starnes 1993 Etheostoma chlorosomum Bluntnose darter Cover, egg attachment Page et al. 1982; Etnier and Starnes 1993 Etheostoma chienense Relict darter Cover, nesting cover, egg Piller and Burr 1999 attachment Etheostoma collis Carolina darker Cover Jenkins and Burkhead 1994 Etheostoma coforosum Coastal darter Cover Mettee et al. 1996 Etheostoma coosae Coosa darter Egg attachment Mettee et al. 1996 Etheostoma corona Crown darter Nesting cover, egg attachment Mettee et aI. 1996 Etheostoma cragini Arkansas darter Cover Pflieger 1997 Etheostoma davisoni Choctawhatchee Cover Mettee et al. 1996 darter Etheostoma gracile Slough darter Cover, egg attachment Braasch and Smith 1967 Etheostoma histrio Harlequin darter Cover, juvenile cover Warren 1982; Pflieger 1997 Etheostoma lachneri Tombigbee darter Cover Mettee et al. 1996 Etheostoma neopterum Lollipop darter Cover, nesting cover, probable Etnier and Starnes 1993 egg attachment Etheostoma nigrum Johnny darter Nesting cover, egg attachment Jenkins and Burhead 1994 Etheostoma olmstedi Tessellated darter Cover, nesting cover, probable Jenkins and Burhead egg attachment 1994 Etheostoma oophylax Guardian darter Cover, nesting cover, probable Etnier and Starnes 1993 egg attachment Etheostonra Goldstripe darter Cover, egg attachment Robison 1977; Johnston parvipinne 1994 Etheosfonru proeliare Cypress darter Cover, egg attachment Burr and Page 1978; Pflieger 1997 Etheostoma pt~nctulatum Stippled darter Cover Pflieger 1997 Etheostonza pyrrhogaster Firebell y darter Probable egg attachment Carney and Burr 1989; Etnier and Starnes 1993 TABLE3. Continued. Species Common name Association with large wood Reference Etheostoma ramseyi Alabama darter Cover Mettee et al. 1996 Efheostoma swaini Gulf darter Cover Etnier and Starnes 1993 Etheostoma tallapoosae Tallapoosa darter Spawning cover, probable Mettee et al. 1996 egg attachment Etheostoma trisella Trispot darter Cover Etnier and Starnes 1993 Etheostoma vitreum Glassy darter Egg attachment Winn and Picciolo 1960 Etheostoma zonale Banded darter Juvenile cover Pflieger 1997 Percina cymatofaenia Bluestripe darter Cover Pflieger 1997 Percina macrocephala Longhead darter Cover Etnier and Starnes 1993 Percina maculata Blackside darter Cover, particularly for Etnier and Starnes 1993; juveniles Pflieger 1997 Percina nigrofasciata Blackbanded darter Cover Etnier and Stames 1993 Percina sciera Dusky darter Cover Page and Smith 1970; Etnier and Starnes 1993; Pflieger 1997 Percina stictogaster Frecklebelly darter Cover Burr and Page 1993; Etnier and Starnes 1993

Dolloff (1995) demonstrated high use by trout of spawning habitat for minnows of the genus habitat created by large wood, they were unable to Cyprinella (Pflieger 1997), which deposit their eggs document a direct relation of trout density to the in crevices. The large range of the blacktail shiner amount of wood in three streams draining wilder- Cyprinella venusta across southeastern U.S. coastal ness areas in the eastem United States. plain streams is thought to be at least partially Unpublished information regarding the role attributable to its ability to use large wood for egg of large wood as habitat for fishes of the Amazon attachment. Madtom catfishes Noturus spp. pro- Basin provides preliminary insights about the role vide extensive care to nests, eggs, and young and of wood in tropical streams (P. Petry, Fish Divi- several species nest on the undersides of large sion, Field Museum of Natural History, Chicago, wood (Burr and Stoeckel1999). The federally en- personal communication). Many species of fishes dangered relict darter attaches its eggs to the un- are believed to use large wood as their habitat in derside of sticks or logs, and the male remains to the Amazon and its . This includes guard the nest site; paucity of woody spawning several groups of catfishes (families Loricariidae, substrates in its sandy coastal plain habitat is a Pseudopimelodidae, Auchenipteridae, and primary factor limiting recruitment of the species Ageneosidae). Species of the family Auchenip- (Piller and Burr 1999).A similar case can be made teridae live among dead trees and branches; they for other egg attaching darters inhabiting coastal are called driftwood catfishes because they are plain and piedmont streams (Table 3). always found inside logs. Several species of elec- tric knifefishes (Order Gymnotiformes) are also frequently found inside logs. In addition to those Beyond Fish fish that actually live inside the logs, many spe- cies are associated with the log jams, including Adding wood to Puerto Rican headwater streams many cichlids and characins. for the purpose of increasing cover for freshwa- ter shrimp did not appear to affect total numbers Large wood can be a vital resource for com- pleting life history phases for a host of fish spe- of shrimp per pool area (Pyron et al. 1999).How- ever, large wood in streams around the world has cies. Almost half the species associations with large wood are related to reproduction or juve- been found to influence a broad array of aquatic nile rearing (Table 3). Fishes living in streams of and riparian dependent organisms in addition to the coastal plain, where streambed materials tend fishes and insects. One of the most spectacular to be fine-grained and highly mobile (Felley 1992), examples of dependence on large wood is the gi- often use large wood for attaching their eggs, ant freshwater crayfish of Tasmania. These ani- mals, known to live for 30 years and reach weights spawning cover, or nest cover. For example, logs with intact, deeply ridged bark provide suitable of 4 kg, depend on wood both for food (they peel FISH RtiLATlONSHII'S WlTII IAHGE WOOD IN SMALL STREAMS 189

and eat layers of rotting eucalyptus logs) and for role of wood in streams including variables such habitat (Hamr 1996). Instream habitat for the Tas- as tree specles, amounts, distribution, relations to manian giant crayfish includes submerged logs, biota (not only fishes but other vertebrates and root masses, and wood accumulations, all of invertebrates), and influence on habitat features which are generally available and abundant in in small streams. Knowledge gaps notwithstand- streams flowing through intact riparian areas. ing, our current knowledge provides a rigorous Another equally unique example is the interac- basis for resource managers to protect, maintain, tion of large wood and the "fishing mussels" and begin the complex task of restoring the func- (Lampsilis australis, L. perovalis, and L. subangulafa) tional character and features of small streams in of the southeastern United States (Haag et al. forested regions.

1995). Females in this group-. release their larvae, which are obligate parasites on fishes, in two dis- crete masses that resemble a pair of small fishes References in shape and color. After release, the lures are teth- Angermeier, P. L., and J. R. Karr. 1984. Relationships ered to the female by a long, transparent mucous between woody debris and fish habitat in a strand. As it undulates in the current, the teth- small warmwater stream. Transactions of the ered lure mimics movement of small fishes (Haag American Fisheries Society 113:716-726. et al. 1995). After some time, the strand detaches Andrus, C. W., B. A. Long, and H. A. Froelich. 1988. from the female, drifts downstream, catches on Woody debris and its contribution to pool for- wood, and continues "fishing" for the host fish mation in a coastal stream 50 years after log- (Micropterus spp.) (Haag et al. 1995; Haag and ging. Canadian Journal of Fisheries and Aquatic Warren 1999). Although the importance of this Sciences 45:2080-2086. "snag and fish" strategy to recruitment has not Aumen, N. G., C. I? Hawkins, and S. V. Gregory. 1990. Influence of woody debris on nutrient retention been documented empirically, the strategy obvi- in catastrophically disturbed streams. Hydrolo- ously extends temporal and spatial exposure of biologia 190:183-192. the lure (and, hence, the larvae) to the target host- Armantrout, N. B., compiler. 1998. Glossary of fish species. aquatic habitat inventory terminology. Ameri- Perhaps less spectacular, but nevertheless can Fisheries Society, Bethesda, Maryland. important, is the association of aquatic turtles and Baker, J. A., and S. T. Ross. 1981. Spatial and tempo- wood. A frequently cited cause of decline of turtles ral resource partitioning by southeastern cyp- in streams and ri;ers is the loss or deliberate re- rinids. Copeia 1981:178-184. moval of large wood from the channel (Felley Becker, G. C. 1983. Fishes of Wisconsin. The Univer- 1992; Buhlmann and Gibbons 1997).Although few sity of Wisconsin Press, Madison. Benda, L., D. Miller, J. Sias, D. Martin, R. Bilby, C. studies have focused on the relationship of turtles Veldhuisen, and T. Du~e2003. Wood recruit- to large wood, snags and logs provide them with ment processes and wood budgeting. Pages 49- basking sites (Conant and Collins 1998), particu- 73 in S. V. Gregory, K. L. Boyer, and A. M. larly the map and false map turtles (genus Gurnell, editors. The ecology and management Graptemys). Other turtle species seem to prefer of wood in world rivers. American Fisheries habitats where wood is present. In Whiskey Chitto Society, Symposium 37, Bethesda, Maryland. Creek, a of the Calcasieu River, Louisi- Benke, A. C., R. L. Henry, 111, D. M. Gillespie, and R. ana, Shively and Jackson (1985) found that the J. Hunter. 1985. Importance of snag habitat for Sabine map turtle was limited to stream reaches production in southeastern streams. with high numbers of snags. The snags provided Fisheries 10:8-13. Benke, A. C., and J. B. Wallace. 2003. Influence of both basking areas and a substrate for algae used wood on invertebrate communities in streams as food by the turtle (Shively and Jackson 1985; and rivers. Pages 149-177 in S. V. Gregory, K. L. Felley 1992). Boyer, and A. M. Gurnell, editors. The ecology In the last 20 years, the list of species associ- and managenlent of large wood in world riv- ated with, if not dependent on, large wood has ers. American Fisheries Society, Symposium 37, grown dramatically. New relationships between Bethesda, Maryland. wood in the water and many taxa of Bilby, R. E., and G. E. Likens. 1980. Importance of emerge regularly, and there clearly is much to be organic debris dams in the structure and func- learned (see also Steel et al. 2003, this volume). tion of stream ecosystems. Ecology 61:1107- Additional research is needed to provide manag- 1113. Bisson, P. A., J. L. Nielson, R. A. Palmason, and L. ers with a more complete understanding of the E. Grove. 1982. A system of naming habitat 190 DOLLOFF AND WARREN

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The Ecology and Management of Wood in World Rivers

Edited by

Stan V. Gregory

Department of Fisheries and Wildlife, Oregon State University Corvallis, Oregon 9733 7, USA

Kathryn L. Boyer

USDA Natural Resources Conservation Service life Habitat Management Institute, Department of Fisheries and Wildlife Oregon State University, Corvallis, Oregon 9733 1, USA

Angela M. Gurnell

Department of Geography, King's College London Strand, London WC2R ZLS, UK

American Fisheries Society Symposium 37

International Conference on Wood in World Rivers eld at Oregon State University, Corvallis, Oregon 23-27 October 2000

American Fisheries Society Bethesda, Maryland 2003

Contents

. . Preface ...... vii Acknowledgments ...... ix Symbols and Abbreviations ...... xi

The Limits of Wood in World Rivers: Present, Past, and Future Ken J. Gregory ...... 1 Geomorphic Effects of Wood in Rivers David R. Montgomery, Brian D. Collins, John M. Buffington, and Timothy B. Abbe ...... Recruitment Processes and Wood Budgeting ee Benda, Daniel Miller, Joan Sias, Douglas Martin, Robert Bilby, rt Veldhuisen, and Thomas Dunne ...... torage and Mobility gela M. Gurnell ...... lic Effects of Wood in Streams and Rivers hael Mutz ...... f Wood in Large Rivers &gay...... on and Nutrient Dynamics of Wood in Streams and Rivers . Bilby ...... od on Invertebrate Communities in Streams and Rivers nke and J. Bruce Wallace ...... with Large Wood in Small Streams loffand Melvin L. Warren, Jr...... s with Wood in Large Rivers ,Magorzata Lapinska, and Peter B. Bayley ...... morphic Effects of Beaver Dams and Their Influence

Morgan Neim, and Danielle Werner ...... nefits of River Wood to Terrestrial and Aquatic

am N. Richards, and Kathryn A. Kelsey ...... v vi CONTENTS Influence of Wood on Aquatic Biodiversity Stezren M. Wondzell nnd Peter A. Bisson ...... 249 Dynamics and Ecological Functions of Wood in Estuarine and Coastal Marine Ecosystems Charles A. Sirnenstad, Alicia Wick, Sfavr Van de Wefering, and Dar~ielL. Botto~n...... 265 Dynamics of Wood in Rivers in the Context of Ecological Disturbance Ft~tosbziNakamura and Frederick J. Swanson ...... 27 Wood in Rivers: A Landscape Perspective Frrderick J. Swanson ...... 29 Modeling the Dynamics of Wood in Streams and Rivers Start V. Gregory, Mark A. Meleason, land Daniel J. Sobota ...... Wood in Streams and Rivers in Developed Landscapes Arturo Elosegi and Lucinda B. Johnson ...... Restoring Streams with Large Wood: A Synthesis Michael Reich, Jeffrey L. Kershner, and Randall C. Wildrnan ...... Wood in River Rehabilitation and Management Timothy B. Abbe, Andrew I? Brooks, and David R. Montgomery ...... Trends in Using Wood to Restore Aquatic Habitats and Fish Communities in Western North American Rivers Peter A. Bissour, Steven M. Wondzell, Gordon H.Reeves, and Stan V: Gregory ...... Riparian Management for Wood in Rivers Kathryn L. Boyer, Dean Rae Berg, and Stan V. Gregory ...... Rivers and Wood: A Human Dimension CropPetts and Robin Welcomme ......