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Changing fish faunas in two reaches of the Rio Grande in the Albuquerque Basin.

Extinctions of native fish and introductions of nonnative fish are changing fish faunas throughout North America (Moyle, 1986; Miller et al., 1989). In the Southwest, degradation of streams and nonnative fishes have caused declines and extinctions among native species (Miller, 1961; Minckley and Deacon, 1968). Entire faunas have been altered dramatically (e.g., Contreras-Balderas et al., 2002; Hoagstrom, 2003), so historical comparisons are necessary to assess ongoing changes.

The native fish fauna of the Rio Grande in the Albuquerque Basin, New Mexico, is in decline and the modern fauna includes many introduced taxa (Platania, 1991). These faunal changes are associated with centuries of development of the watershed by humans, which intensified after 1846 (Horgan, 1984; Kelley, 1986; Scurlock, 1998). Presently, waters of the Rio Grande are heavily exploited and tightly controlled (Price et al., 2007). Our goals in this paper are to characterize the modern fish fauna of two reaches of the Rio Grande in the Albuquerque Basin, assess short-term faunal change, and consider long-term faunal changes with regard to chronology of settlement by humans and extirpations of native fish.

MATERIALS AND METHODS--Study Area--With the exception of its mountainous headwaters, the Rio Grande upstream of the Rio Conchos (a major tributary in chihuahua) traverses a rift valley constructed via tectonic, volcanic, and fluvial processes (Bryan, 1938; Hawley, 1978). The river flows through a sequence of structural basins separated by narrows (Lee, 1907; Bryan, 1938). Basins provide ideal conditions for irrigated agriculture and urban development, whereas narrows provide ideal locations for dams that divert river water onto basins downstream (Blaney et al., 1938; Towne, 2007).

The Albuquerque Basin lies in the central segment of the Rio Grande Rift in central New Mexico (Chapin, 1979). It extends ca. 164 km north to south, 40-64 km east to west, and is filled to a depth of 3,658 m with sandstone, mudstone, and gravel (Kelley, 1977). The pre-development floodplain of the Rio Grande in the Albuquerque Basin was 3-5 km in width, except for narrows near san Felipe (91 m in width) and Isleta (<2 km in width; Kelley, 1977). These narrows divide the Albuquerque Basin into (from upstream to downstream) the Santo Domingo, Albuquerque, and Belen valleys (Kelley, 1977). Diversion dams were constructed at each narrows (Angostura Dam near san Felipe, 1936; Isleta Dam in the Isleta narrows, 1936) to irrigate valleys downstream. Diversion dams also were placed at the head of the Santo Domingo Valley (Cochiti Diversion Dam, 1936) and at the mouth of the Belen Valley (san Acacia Diversion Dam, 1936). The four dams combined to fragment the Rio Grande of the Albuquerque Basin into three discreet reaches that corresponded to the three valleys.

Agricultural and urban development substantially modified the Rio Grande (Stafford, 1938; Woodson and Martin, 1965; Schmidt et al., 2003) and, unfortunately, pre-development conditions were never quantified. A voyage from Santo Domingo to Mesilla, New Mexico, in August 1874 suggested the river was navigable despite substantial withdrawals of water (Kelley, 1986), but 1874 was a high-water year (Scurlock, 1998) and may not represent average pre-settlement conditions. Nevertheless, in that year, the only specimens of the shovelnose sturgeon (Scaphirhynchus platorynchus) ever collected from the Rio Grande were taken at Albuquerque (Cope and Yarrow, 1875), perhaps signaling the end of an era because development proceeded rapidly throughout the late 19th century and withdrawals for irrigation largely depleted the Rio Grande by 1900 (Kelley, 1986; Scurlock, 1998).

The historical channel of the Rio Grande in the Albuquerque Basin was relatively wide and braided (Stafford, 1938; Scurlock, 1998). With the exception of the narrows near san Felipe, it was a sand-bed channel (Culbertson and Dawdy, 1964). Sands originated from the Rio Jemez, Rio Grande upstream, Galisteo Creek, and Santa Fe River (Rittenhouse, 1944), but flood-control dams (Jemez Canyon Dam, 1953; Galisteo Dam, 1970; Cochiti Dam, which replaced Cochiti Diversion Dam, 1973) now capture sand inflows from each of these sources. Deprivation of sediment has caused geomorphic degradation, including narrowing of the river channel and coarsening of the substrate throughout the Albuquerque Basin, most severely in proximity to the sediment-capturing dams (i.e., in the Santo Domingo and Albuquerque valleys), but effects increasingly extend downstream (Dewey et al., 1979; Lagasse, 1994; Schmidt et al., 2003).

Our study included the Rio Grande in the Albuquerque (61 river km) and Belen (90 river km) valleys of the Albuquerque Basin. Flow regimes in both reaches largely reflected patterns of release from Cochiti Dam, but flows within the Belen Valley were less than in the Albuquerque Valley during spring and summer because of withdrawals made via the Isleta Dam (Fig. 1a), which separates the two reaches. The Albuquerque Valley also had a coarser-sand substrate than the Belen Valley (Figs. 1b and c), which reflected the proximity of sediment-capturing dams (Lagasse, 1994; Schmidt et al., 2003).

Collections--We collected fishes with flat seines (3.2-mm mesh, 3.0-m wide, 1.2-m deep; one lead weight every 15 cm). Sampling trips were designed to determine the fish fauna of each reach, so sampling locations were visited throughout the length of each reach during each trip. Within sampling locations, distinct mesohabitats were seined in proportion to their presence. Each seine collection was made within a single mesohabitat type, including riffles, runs, submerged banks, pools, plunge pools, channel-confluence pools, embayments, forewaters, backwaters, and isolated pools. Piles of wood and associated materials created habitat in the river channel and, when present, were sampled. Distance seined was measured for each collection and multiplied by width of seine to quantify the area sampled (sampling effort). In most cases, fishes were identified and enumerated in the field immediately upon collection and released alive. Data from collections of fish were recorded separately, by seine collection. In some cases, voucher specimens were retained and deposited in the Division of Fishes, Museum of Southwestern Biology, University of New Mexico, Albuquerque (MSB-ACC2003-X:1).

[FIGURE 1 OMITTED]

Comparisons of Reaches--We contrasted abundance of fish between reaches by comparing mean density (Mann-Whitney test) and percentage of seine hauls with no fish. We investigated both structure of the assemblage of fish faunas and faunal similarity between valleys. Marsh-Matthews and Matthews (2000) suggested the two may be incongruent across space and time. We quantified structure of assemblage by determining taxon richness (number of taxa collected) and constructing taxon-abundance distributions, which simultaneously indicate taxonomic richness (number of species present) and patterns of dominance of taxa (species that are most abundant; Tokeshi, 1993). We compared taxon-abundance distributions with a two-sample Kolmogorov-Smirnoff test (Tokeshi, 1993). We also used Fisher's [alpha] (Magurran, 2003) to estimate taxonomic diversity within each valley. Fisher's [alpha] is a robust measure of diversity that is not biased by highly abundant taxa, rare taxa, or sample size (Kempton and Taylor, 1974). Also, confidence limits can be calculated and used to statistically compare diversity among faunas (Magurran, 2003). To compare faunal similarity between valleys, we used Morisita's Index ([I.sub.m]), which is accurate (Smith and Zaret, 1982) and minimally biased by sample size or taxonomic diversity (Wolda, 1981). index values range from zero to slightly greater than one, with those >0.7 being considered relatively strong (Ross et al., 1985).

We designated taxa as native or nonnative and constructed an overall faunal list of fishes for each valley based on data provided by Miller (1977), Smith and Miller (1986), Sublette et al. (1990), Platania (1991), Rinne and Platania (1995), Propst (1999), and Calamusso et al. (2005). We consulted multiple sources because there is disagreement on the status of some fishes (Table 1). Nonnative taxa were further classified as either drainage nonnatives (native to the Rio Grande drainage but not the study area) or extra-drainage nonnatives (not native anywhere in the Rio Grande drainage). Drainage nonnatives potentially could invade the study area via their own means (although it is unlikely they could disperse upstream past major dams) and are more likely to successfully colonize the study area because they naturally inhabit similar environments (Gido et al., 2004). In contrast, extra-drainage nonnatives could only reach the study area with aid of humans and are less likely on average to successfully invade because they originate in more distant habitats with less similar environmental conditions.

Historical Comparisons--We assessed short-term change in faunas of both valleys by comparing our findings with those of Platania (1991). He presented data from collections made during August 1984 at six locations in the Albuquerque Valley and three in the Belen Valley. We compared taxon diversity, taxon-abundance distributions, faunal similarity, and prevalence of nonnatives between his and our study. We also assessed long-term change in the combined fish fauna by comparing the number of native and nonnative taxa that have been extirpated from the study area with the number present in our study.

RESULTS--Collecting Effort and Abundance of Fish--We made collections during September 1998-April 2001, conducting 5,570 seine collections in 28 sampling trips within the Albuquerque Valley and 4,692 seine collections in 19 sampling trips within the Belen Valley. Total area seined was 128,980 [m.sup.2] in the Albuquerque Valley and 99,415 [m.sup.2] in the Belen Valley. Collections were made in every month of the year in both reaches. Mean density of fish was significantly different between reaches of the river (Mann-Whitney U = 9,572,604, P < 0.001), being nearly double in the Belen Valley (1.1 [+ or -] 5.74 SD fish/ [m.sup.2]) compared to the Albuquerque Valley (0.6 [+ or -] 4.24 SD fish/[m.sup.2]). Fishes were absent from 63% of all seine collections in the Albuquerque Valley compared to 40% in the Belen Valley.

Faunal Diversity and Similarity--Most fishes were present in both reaches of the river. Exceptions were the extra-drainage nonnatives brown trout (Salmo trutta) and yellow perch (Perca flavescens) that were collected only from the Albuquerque Valley, and the drainage nonnative largemouth bass (Micropterus salmoides) that was collected only from the Belen Valley (Table 1). Hence, taxon richness was similar (20 fishes in the Albuquerque Valley, 19 in the Belen Valley), as was taxon diversity (Fig. 2d). Taxon-abundance distributions indicated similar faunal structure between valleys (Figs. 3c and 3d; Z = 0.53, P = 0.945). However, faunal similarity was moderate ([I.sub.m] = 0.6), indicating that taxa varied in relative dominance between valleys. For instance, drainage nonnatives dominated the Albuquerque Valley, whereas natives dominated the Belen Valley (Fig. 2b).

Short-term Historical Change--We collected more taxa from the Albuquerque Valley in 1998-2001 than were collected in 1984 (20 versus 17), but taxonomic diversity was higher in 1984 (Figs. 2c and 2d) because dominance was more even among taxa (Figs. 3a and 3c). The native Rio Grande chub (Gila pandora) and the extra-drainage nonnative black bullhead (Ameiurus melas) were present in 1984 but missing in 1998-2001, whereas the native gizzard shad (Dorosoma cepedianum) and extra-drainage nonnative brown trout, white bass (Morone chrysops), spotted bass (Micropterus punctulatus), yellow perch, and walleye (Sander vitreus) were present in 1998-2001, but missing in 1984. Taxon-abundance distributions indicated similar structure of assemblages between faunas (Figs. 3a and 3c; Z = 0.86, P = 0.456), but faunal similarity was relatively low ([I.sub.m] = 0.4), indicating substantial change in dominance of taxa, i.e., drainage nonnatives were more dominant in 1998-2001 than in 1984 (Figs. 2a and 2b). Notably, the white sucker (Catostomus commersonii) and western mosquitofish (Gambusia affinis) increased, whereas the native Rio Grande silvery minnow (Hybognathus amarus) and longnose dace (Rhinichthys cataractae) declined (Table 1, Figs. 3a and 3c). The native red shiner (Cyprinella lutrensis) was dominant in both faunas and, aside from gizzard shad, which was rare in 1998-2001 but absent in 1984, the red shiner was the only native taxon that did not decline in the Albuquerque Valley between periods (Table 1).

Taxonomic richness in the Belen Valley differed between 1984 and 1998-2001 (19 versus 13 taxa), but diversity was similar (Figs. 2c and 2d). We collected all taxa present in 1984 plus the native gizzard shad, the drainage nonnatives green sunfish (Lepomis cyanellus) and bluegill (Lepomis macrochirus), and the extra-drainage nonnatives white bass, spotted bass, and walleye. Taxon-abundance distributions indicated similar structure of assemblages between faunas (Figs. 3b and 3d; Z = 1.04, P = 0.235) and faunal similarity was high ([I.sub.m] = 0.9), indicating minimal change in relative dominance of taxa. However, drainage nonnatives were more dominant in 1998-2001 than in 1984 (Figs. 2a and 2b). The most dramatic faunal change in the Belen Valley was decline of the native Rio Grande silvery minnow, which was offset by increasing dominance of the native red shiner and drainage nonnative western mosquitofish (Table 1, Figs. 3b and 3d). In contrast to the Albuquerque Valley, several other native taxa (fathead minnow Pimephales promelas, flathead chub Platygobio gracilis, slender carpsucker Carpiodes carpio) also increased in dominance between 1984 and 1998-2001 (Table 1). However, their increases were small compared to the red shiner (< 4%), and the flathead chub was rare in both periods (<0.4%).

Long-term Historical Change--Between 1998 and 2001, we collected 21 of the 44 fishes previously reported from the Albuquerque and Belen valleys or presumed native there (Table 1). This indicates substantial long-term faunal change. Drainage nonnatives showed highest persistence as a group, with six of eight documented taxa (75%) remaining. In contrast, only 6 of 14 known extra-drainage nonnatives (43%) were present in 1998-2001 and only 7 of 22 natives (32%) remained. Only one of the missing natives (Rio Grande chub) and one of the missing extra-drainage nonnatives (black bullhead) were present in 1984, indicating that most extirpations and failed introductions occurred prior to then. To our knowledge, this is the first report of spotted bass from the mainstem Rio Grande.

DISCUSSION--Comparisons of Reaches and Short-term Historical Change--Marsh-Matthews and Matthews (2000) suggested properties of structure of assemblages (i.e., taxon richness, taxon-abundance distributions) are relatively stable at a regional scale, whereas faunal composition (relative dominance) depends on local conditions. Our study supports their hypothesis. Taxon richness and taxon-abundance distributions in the Albuquerque Basin were stable between reaches of the river and study periods although sensitive, native taxa declined and nonnatives were introduced and proliferated.

[FIGURE 2 OMITTED]

Fish faunas of the two valleys showed different trends between 1984 and 1998-2001. Substantial change occurred in the Albuquerque Valley, with dramatic declines of sensitive natives and concomitant increases of nonnatives. The Albuquerque Valley is apparently becoming less suitable for native fishes, with the exception of the red shiner, which has high environmental tolerances (Matthews, 1985, 1987) and often is invasive when introduced into altered habitats outside its native range (Douglas et al., 1994; Propst and Gido, 2004). Faunal change was less dramatic in the Belen Valley, which appeared to remain relatively suitable for three tolerant natives: red shiner, fathead minnow, and slender carpsucker. Thus, although structure of assemblages remained similar between valleys and time periods, faunal similarity diverged, presumably reflecting divergence in environmental conditions (e.g., discharge in spring and summer, coarseness of substrate).

Remaining native fishes are presumably the hardiest and most suited to live in human-altered habitats. Nevertheless, the Rio Grande silvery minnow and longnose dace appear to be succumbing to ongoing environmental degradation. The Rio Grande silvery minnow, a federally listed endangered species, declined rapidly after Cochiti Dam was completed (Bestgen and Platania, 1991); perhaps, in response to oligotrophication and loss of floodplain connectivity (Cowley et al., 2006; Shirey et al., 2008). Displacement of pelagic embryos and larvae through diversion dams also limits recruitment in situ and degrades genetic diversity (Turner et al., 2006; Dudley and Platania, 2007). Continued decline of this taxon has led to intensifying conservation and recovery efforts (Parody, 2007; Towne, 2007).

[FIGURE 3 OMITTED]

Recent decline of the longnose dace has not been studied, but is unlikely to be caused by downstream displacement because embryos and larvae of this taxon are not pelagic (Sublette et al., 1990). Dewatering may contribute to decline of the longnose dace because it usually is associated with relatively swift and cool waters (Koster, 1957; Sublette et al., 1990). Reductions in streamflow typically reduce velocities of water and increase diel fluctuations in temperature.

Absence of some fishes from our collections might be an artifact of sampling bias. Fishes that occur upstream in the Rio Grande or in adjacent waters, such as floodplain wetlands, artificial ponds and lakes, or canals, might be transient within the Rio Grande, present only under suitable conditions, or when displaced by high flows. For example, the native Rio Grande chub, as well as the nonnative central stoneroller (Campostoma anomalum), black bullhead, and rainbow trout (Oncorhynchus mykiss), are present in adjacent canals (Platania, 1991; Cowley et al., 2007) and might, at times, inhabit the mainstem Rio Grande. Some taxa present in canals and other adjacent waters, such as the rainwater killifish (Lucania parva) and longear sunfish (Lepomis megalotis; Sublette et al., 1990; Platania, 1991; Cowley et al., 2007) have yet to be reported from the mainstem Rio Grande, but could invade if conditions become suitable.

Fishes of all kinds had low abundance in the study area, particularly in the Albuquerque Valley. Seine collections associated with shorelines, woody cover, slow velocity, and shallow depth often produced fishes, whereas seine collections from fluvial habitats frequently produced none. Conditions apparently have become unsuitable for fluvial fishes (those that occupy flowing-water habitats); perhaps, because periods of intermittent streamflow preclude successful reproduction (Durham and Wilde, 2006) or favor taxa that tolerate harsh conditions in isolated pools (Ostrand and Wilde, 2004). only three native fluvial fishes remain (Rio Grande silvery minnow, flathead chub, longnose dace). Of these, the flathead chub is uncommon and the other two taxa are in decline.

Long-term Historical Change--our estimate of 22 native fishes within the Albuquerque and Belen valleys is conservative because it is based on scant historical and archaeological evidence. Although presently fragmented, the Rio Grande was once a free-flowing river of substantial size throughout its length (Horgan, 1984; Kelley, 1986). It is unclear what, if any, barrier would have precluded fishes presently restricted to the lower Rio Grande from ascending upstream to the Albuquerque and Belen valleys, especially during high-flow periods. Indeed, historical presence of catadromous American eels (Anguilla rostrata) indicates such a migration was possible (Koster, 1957). Riverine fishes that were never recorded from the Albuquerque and Belen valleys but potentially ranged upstream prior to construction of dams and desiccation of the river include the alligator gar (Atractosteus spatula), spotted gar (Lepisosteus oculatus), ghost shiner (Notropis buchanani), black buffalo (Ictiobus niger), and Mexican redhorse (Moxostoma austrinum).

Extirpations of native fish from the Albuquerque and Belen valleys occurred in two periods and primarily involved fluvial fishes. The first extirpation period (pre-1915) resulted in disappearance of large-bodied, big-river natives (shovelnose sturgeon, longnose gar Lepisosteus osseus, American eel, blue sucker Cycleptus elongatus, smallmouth buffalo Ictiobus bubalus, gray redhorse Moxostoma congestum, blue catfish Ictalurus furcatus, flathead catfish Pylodictis olivaris, freshwater drum Aplodinotus grunniens) that were only known from archaeological evidence or 19th-century surveys. Due to scant evidence, it is not certain when extirpations began, but all taxa appear to have been present in 1450 and some were present as late as 1874 (Sublette et al., 1990). This period of extirpation was associated with increasing aridity and human-caused desiccation of the Rio Grande (Kelley, 1986; Smith and Miller, 1986; Scurlock, 1998). If any big-river fishes persisted until 1915, completion of Elephant Butte Dam likely finalized their disappearance by blocking dispersal and isolating upstream populations.

The second extirpation period (post-1939) resulted in disappearance of native fluvial minnows (Rio Grande speckled chub Macrhybopsis aestivalis, Rio Grande shiner Notropis jemezanus, phantom shiner Notropis orca, Rio Grande bluntnose shiner Notropis simus) that persisted long enough to be documented by historical surveys of the mid-20th century. These taxa disappeared between 1939 and 1988 (Chernoff et al., 1982; Bestgen and Platania, 1990). Their extirpations were facilitated by increasing aridity and human-caused dewatering that caused earlier extirpations of big-river fishes, but the fluvial minnows withstood periodic dewatering; perhaps, by retreating to more-permanent reaches of the river, such as those in narrows and canyons (sensu Minckley, 1991). However, intensifying effects of building dams, manipulation of the river channel, urbanization, and invasions of nonnative organisms along with increasing withdrawals of water in the 20th century ultimately corresponded with extirpations of fluvial minnows (Bestgen and Platania, 1990, 1991; Scurlock, 1998). Not all native fluvial minnows have been extirpated, but rarity of the flathead chub and recent decline of the Rio Grande silvery minnow and longnose dace suggest they could be.

Synergism of increasing dominance by nonnative fishes and changing environmental conditions seems most likely to cause future extirpations of native fishes in the Albuquerque and Belen valleys. For instance, some nonnatives (common carp Cyprinus carpio, white sucker) spawn earlier than natives, which could give them a competitive advantage because larval fishes of all taxa rely on the same nursery habitats and the same types of prey, especially when the Rio Grande dries (Pease et al., 2006). Also, carnivorous nonnatives that inhabit canals might prey upon natives because when the Rio Grande dries, riverine fishes retreat to canals as refuges (Cowley et al., 2007). These interactions likely create a combined stressor on natives.

The fish fauna of the Rio Grande in the Albuquerque and Belen valleys is changing. Long-term changes (decades to centuries) have been dramatic, including disappearance of most native fishes. Short-term changes (years to decades) have been more subtle, involving changing patterns of dominance among fishes. it is uncertain whether changing dominance portends future extirpations or simply reflects short-term fluctuations in populations. If it indicates long-term trends, then relict populations of sensitive, native fishes appear to be threatened with extirpation.

This project was partially supported by the United States Army Corps of Engineers and the city of Albuquerque. L. Hall prepared the resumen. Two anonymous reviewers made improvements to the manuscript. Field assistance was provided by A. M. Andazola, C. S. Berkhouse, J. R. Brooks, S. J. Bulgrin, C. A. Cave, S. R. Davenport, J. E. Davis, G. B. Dean, J. S. Donahue, R. A. Fike, M. Harberg, P. A. Hoban, A. L. Hobbes, J. A. Jackson, J. S. Johns, J. Kimball, R. D. Larson, D. Lente, W. C. Liebfried, B. D. Lujan, M. V. McPhee, M. L. Meneks, M. D. Morens, A. R. Nevarez, B. Ortiz, S. D. Parris, J. W. Sigler, H. E. Watts, and B. G. Wiley.

Submitted 26 August 2008. Accepted 1 June 2009. Associate Editor was Gary P. Garrett.

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CHRISTOPHER W. HOAGSTROM, * W. JASON REMSHARDT, JUDE R. SMITH, AND JAMES E. BROOKS

New Mexico Fish and Wildlife Conservation Office, United States Fish and Wildlife Service, 3800 Commons Avenue NE, Albuquerque, NM 87109

Present address of CWH: Department of Zoology, Weber State University, 2505 University Circle, Ogden, UT 84408

Present address of JRS: Buffalo Lake National Wildlife Refuge, P.O. Box 179, Umbarger, TX 79091

* Correspondent: ChristopherHoagstrom@weber.edu
TABLE 1--Fishes of the Albuquerque and Belen valleys, Albuquerque
Basin, Sandoval, Bernalillo, Valencia, and Socorro counties, New
Mexico, including all taxa reported from the study area or presumed
native based on archaeological evidence. Dominance (percentage
abundance) is indicated for taxa collected in 1984 (Platania, 1991)
and during 1998-2001 (this study).

                                            Albuquerque Valley

         Fish taxa            Status (a)      1984      1998-2001

Scaphirhynchus platorynchus   Native           --          --
Lepisosteus osseus            Native (b)       --          --
Anguilla rostrata             Native           --          --
Dorosoma cepedianum           Native (c)       --          0.01
Campostoma anomalum           D.nonn.          --          --
Cyprinella lutrensis          Native          11.62       28.38
Cyprinus carpio               E.nonn.          0.22        0.36
Dionda episcopa               D.nonn. (d)      --          --
Gila pandora                  Native           0.04        --
Hybognathus amarus            Native          24.10        0.24
Macrhybopsis aestivalis       Native           --          --
Notemigonus crysoleucas       E.nonn.          --          --
Notropis jemezanus            Native           --          --
Notropis orca                 Native           --          --
Notropis simus                Native           --          --
Pimephales promelas           Native           5.10        1.49
Platygobio gracilis           Native           1.77        1.49
Rhinichthys cataractae        Native          23.11        1.14
Carpiodes carpio              Native           4.67        3.20
Catostomus commersonii        D.nonn.          3.89       41.41
Catostomus plebeius           Native           --          --
Cycleptus elongatus           Native (b)       --          --
Ictiobus bubalus              Native (b)       --          --
Moxostoma congestum           Native (b)       --          --
Ameiurus melas                E.nonn. (e)      0.91        --
Ameiurus natalis              E.nonn.          4.45        0.19
Ictalurus furcatus            Native           --          --
Ictalurus punctatus           D.nonn.          7.78        3.42
Pylodictis olivaris           Native (b)       --          --
Oncorhynchus mykiss           E.nonn.          --          --
Salmo trutta                  E.nonn.          --          0.01
Gambusia affinis              D.nonn. (f)     11.10       18.19
Morone chrysops               E.nonn.          --          0.01
Lepomis cyanellus             D.nonn.          0.26        0.06
Lepomis gulosus               E.nonn. (g)      --          --
Lepomis macrochirus           D.nonn. (h)      0.04        0.03
Micropterus dolomieu          E.nonn.          --          --
Micropterus salmoides         D.nonn.          0.04        --
Micropterus punctulatus       E.nonn.          --          0.02
Pomoxis annularis             E.nonn.          0.91        0.33
Pomoxis nigromaculatus        E.nonn.          --          --
Perca flavescens              E.nonn.          --          0.01
Sander vitreus vitreus        E.nonn.          --          0.02
Aplodinotus grunniens         Native (b)       --          --

                                   Belen Valley

         Fish taxa               1984       1998-2001

Scaphirhynchus platorynchus       --           --
Lepisosteus osseus                --           --
Anguilla rostrata                 --           --
Dorosoma cepedianum               --           0.01
Campostoma anomalum               --           --
Cyprinella lutrensis             52.64        60.83
Cyprinus carpio                   0.30         1.20
Dionda episcopa                   --           --
Gila pandora                      --           --
Hybognathus amarus               30.21         0.17
Macrhybopsis aestivalis           --           --
Notemigonus crysoleucas           --           --
Notropis jemezanus                --           --
Notropis orca                     --           --
Notropis simus                    --           --
Pimephales promelas               1.72         5.64
Platygobio gracilis               0.15         0.38
Rhinichthys cataractae            0.07         0.01
Carpiodes carpio                  7.94        10.38
Catostomus commersonii            0.97         1.07
Catostomus plebeius               --           --
Cycleptus elongatus               --           --
Ictiobus bubalus                  --           --
Moxostoma congestum               --           --
Ameiurus melas                    --           --
Ameiurus natalis                  1.42         0.35
Ictalurus furcatus                --           --
Ictalurus punctatus               2.55         4.04
Pylodictis olivaris               --           --
Oncorhynchus mykiss               --           --
Salmo trutta                      --           --
Gambusia affinis                  1.91        15.39
Morone chrysops                   --          <0.01
Lepomis cyanellus                 --          <0.01
Lepomis gulosus                   --           --
Lepomis macrochirus               --          <0.01
Micropterus dolomieu              --           --
Micropterus salmoides             0.07         0.42
Micropterus punctulatus           --           0.01
Pomoxis annularis                 0.04         0.08
Pomoxis nigromaculatus            --           --
Perca flavescens                  --           --
Sander vitreus vitreus            --           0.01
Aplodinotus grunniens             --           --

(a) Categories of status of taxa are: Native, native to the study
area; D.nonn., native to the Rio Grande drainage, but not the
Albuquerque Basin (drainage nonnative); and E.nonn., nonnative to
the entire Rio Grande drainage (extra-drainage nonnative).

(b) Taxa presumed native based on archaeological evidence reviewed
by Sublette et al. (1990).

(c) Authors disagree about the status of D. cepedianum. We follow
Miller (1977), Sublette et al. (1990), Propst (1999), and Calamusso
et al. (2005).

(d) Authors disagree about the status of D. episcopa. We follow
Smith and Miller (1986), Rinne and Platania (1995), Propst (1999),
and Calamusso et al. (2005).

(e) Some authors list A. melas as native elsewhere in the Rio
Grande drainage (Smith and Miller, 1986; Edwards et al., 2002). We
follow Miller (1977) and Calamusso et al. (2005).

(f) Authors disagree about the status of G. affinis. We follow
Miller (1977), Smith and Miller (1986), Platania (1991), Rinne and
Platania (1995), and Calamusso et al. (2005).

(g) Some authors list L. gulosus as native elsewhere in the Rio
Grande drainage (Miller, 1977; Smith and Miller, 1986; Edwards et
al., 2002). We follow Sublette et al. (1990), Contreras-Balderas et
al. (2002), and Calamusso et al. (2005).

(h) Authors disagree about the status of L. macrochirus. We follow
Miller (1977), Smith and Miller (1986), Platania (1991), Rinne and
Platania (1995), and Calamusso et al. (2005).
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Author:Hoagstrom, Christopher W.; Remshardt, W. Jason; Smith, Jude R.; Brooks, James E.
Publication:Southwestern Naturalist
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Date:Mar 1, 2010
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