Selected aspects of the natural history of the desert sucker ( (Pantosteus) clarkii)

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Selected aspects of the natural history of the desert sucker [Catostomus (Pantosteus) clarkii]

Ivanyi, Craig Stephen, M.S.

The University of Arizona, 1989

UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106

Selected Aspects of the Natural History

of the Desert Sucker

r Catostomus (Pantosteus) clarkiil

by

Craig Stephen Ivanyi

A Thesis Submitted to the Faculty of the

SCHOOL OF RENEWABLE NATURAL RESOURCES

In Partial Fulfillment of the Requirements

For the Degree of

MASTERS OF SCIENCE

WITH A MAJOR IN WILDLIFE AND FISHERIES SCIENCE

In the Graduate College

THE UNIVERSITY OF ARIZONA

19 8 9 STATEMENT BY AUTHOR 2 This thesis has been submitted in partial fulfillment of requirements for an advanced degree at The University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the Library.

Brief quotations from this thesis are allowable without special permission, provided that accurate acknowledgment of the source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the head of the major department or the Dean of the Graduate College when in his or her judgment the proposed use of the material is in the interests of scholarship. In all other instances, however, permission must be obtained from the author.

APPROVAL BY THESIS COMMITTEE

on the date shown below:

rector Assistant Unit Leader, Arizona Cooperative Fish and Wildlife Research Unit

0. Eugene Maughan Unit Leader, Arizona Cooperative Fish and Wildlife R ? William J. MatterJL Associate Professorofe of Fisheries Science ACKNOWLEDGEMENTS 3

I wish to express my gratitude to Charles D. ziebell for his guidance and suggestions throughout this study. I also thank William Hatter and Gene Maughan for their help in the development of this project and preparation of this manuscript. My thanks also go to Carol Ivanyi, Hugh

McCrystal, and Lynn Ramsey for their ideas, field assistance as well as much needed mental support. Many others "assisted with aspects of this study, and to them I owe many thanks; Field assistance: Cord Ivanyi, Marcy

Owens, Don Davison, Laura Duckett, Rick Roy; Art and photography: Kelly Ivanyi and Ken Wintin; Advice: James

Jarchow D.V.M. and Jeff Hill.

This project was supported by the Arizona Cooperative

Fish and Wildlife Research Unit, which is maintained by The

University of Arizona, the Arizona Game and Fish

Department, and the U. S. Fish and Wildlife Service. TABLE OF CONTENTS 4

Page

LIST OF FIGURES 5

ABSTRACT 6

INTRODUCTION 7

DESCRIPTION OF STUDY AREAS 10

METHODS 13

RESULTS 15

DISCUSSION 21

CONCLUSIONS/MANAGEMENT IMPLICATIONS 26

APPENDIX A: Sonoita Creek Pool Description 29

LITERATURE CITED 30 5

LIST OF FIGURES

Figure Page

1. Length-Weight Relationship Curve 16

2. Length Frequency Histogram 18

3. Catostomus (Pantosteus) clarkii Color Phases..19 ABSTRACT 6

Selected aspects of the life history of desert suckers

[CatogtQimAS (Pantosteus) clarkiil were examined in four southeastern Arizona streams. Diet, age class structure, reproduction, and habitat use were analyzed from field data obtained from April 1987 through October 1988. Adults fed primarily on plant matter, while fry fed almost exclusively on . Aging of suckers by scales was not possible and length frequencies were too evenly distributed to determine age class structure. In 1988 ovary and teste development occurred from late January through April, with spawning commencing in May. Suckers primarily used pools with high water flow and significant (~80%) cover formed by tree limbs, branches, leaves, and other debris.

Deterioration or loss of suitable habitat is reducing the range of the sucker due to reductions in and manipulations of surface waters. INTRODUCTION 7

Introduction of exotic species and manipulation of available waters through impoundments, diversion and pumping of underground aquifers, have caused many of

Arizona's native ichthyofauna to face a tenuous existence

(Hastings and Turner, 1965; Minckley, 1973). Some fish species are extinct and many are endangered (Naiman and

Soltz, 1981). Currently 25 of 35 native species are listed by the Arizona Game and Fish Dept. as threatened, 15 of these are federally listed as threatened or endangered

(Hendrickson and Simons, 1988). Many of the smaller rivers and creeks of Arizona are now dry except during periods of high precipitation or snowmelt (Minckley, 1973). As a result dry, shifting washes with little or no fish habitat have replaced lush cienegas where fish were abundant

(Hendrickson and Simons, 1988). According to the United

States Dept. of the Interior, Bureau of Reclamation (1975) human demands for water do, and will continue to, exceed current supplies, and that demands are increasing as

Arizona's population continues to grow. Water depletion is certain to put additional pressure on Arizona's remaining native fishes. Our poor knowledge of specific biological requirements of these species further increase the chance of irreparable damage to native fish populations.

One of these native fishes is Catostomus (Pantosteus) 8 clarkii^. known commonly as the desert sucker (Eddy and

Underhill, 1978). This sucker is found in the Colorado

River drainage below the Grand Canyon, the system of Arizona and New Mexico, and the Virgin and the

White Rivers of Utah and Nevada (Eddy and Underhill, 1978).

This fish attains standard lengths up to 325 mm and can be identified by its subterminal mouth with large lips and small papillae on the lower lip and oral face of the upper lip. The sucker possesses a cartilaginous sheath inside the mouth on upper and lower jaws, which is used for scraping, and a darkened peritoneum, both considered to be adaptations associated with an herbivorous lifestyle

(Smith, 1966; Smith and Koehn, 1971).

A limited amount of information is currently available on the natural history of the sucker. Mpoame (1982) and

Wier et al. (1983) examined its parasites in Aravaipa

Creek, Arizona. Lowe et al. (1967) studied tolerance to low oxygen tensions for the sucker and Siebert (1980),

Bestgen et al. (Unpubl. data) and Meffe (1987) documented movements in Arizona and New Mexico streams. Several authors have examined its distribution (Barber and

Minckley, 1966; Kepner, 1979; Armantrout, 1983; Deacon,

1983; McNatt, 1983; Cross, 1985). Fisher et al. (1981)

1 1I adopt the "ii" ending, for £_(£i.) clarkii. in accordance to the International Code of Zoological Nomenclature (1964), article 32, page 35, letter "a", regarding "original spelling" of scientific names. 9 found the sucker to be herbivorous, feeding heavily on epipelic diatoms in Sycamore Creek, Arizona. They also noted crepuscular peaks in the sucker's diel feeding behavior.

Despite these previous studies, much remains to be learned about the life history of the sucker. The purpose of this study was to obtain information on diet, age class structure, length-weight relationships, reproduction and fecundity, and habitat use. This additional data will provide a better understanding of the species. DESCRIPTION OF STUDY AREAS 10

Four streams in southeast Arizona were studied.

Limited data were collected from O'Donnell Creek, the

Babocomari River and Bass , all tributaries of

the San Pedro River. Most of the data were collected from

Sonoita Creek, a tributary of the Santa Cruz River. These

streams are all relatively small first or second order

streams, with riparian zones dominated by cottonwoods

(populug §£.), willows f Salix se. ), ashes (FraxiBMs SS>.), and to a lesser degree walnuts (Jualans sp.) and sycamores

(Platanus sp.). O'Donnell Creek consisted of shallow

(7.5-15.0 cm deep), narrow (<1 m) riffles and pools of

various sizes (most <0.5 m deep and 1-2 m in diameter).

Slow-moving water traversed smooth bedrock and fine sand

and silt substrates. The creek flows northeast from Canelo

Hills to the (T21S, R18E; to T21S, R19E).

The sample site was at an elevation of 1600 m. Thick

streamside vegetation made up of grasses, rushes and

horsetail (Equisetum sp.) was prevalent.

The Babocomari River originates at an elevation of

-1500 m in a grassy cienega and flows east from near Elgin to Fairbank (T20S, R18E to T20S, R21E). This very slow

moving stream was characterized by small riffles (7-15 cm deep, <0.5 m wide) and small almost motionless pools (0.25-

1.0 m deep, 1-2 m in diameter). A few pools were 3-4 m deep and 5 m in diameter. Substrates consisted 11

mostly of silty/muddy sands and some small boulder/cobble

areas. A man-made dam near the Babocomari Ranch House,

forms a pond approximately 20 m in diameter that was

surrounded by rushes and sedges.

Bass Canyon Creek runs south from the Galiuro

Mountains to the Muleshoe Ranch (T11S, R21E to T12S, R21E)

where it joins Hot Springs Creek at an elevation of 1300 m.

Substrates included cobbles and boulders with some sandy

bottomed pools. Riffles varied from 3 cm-0.5 m deep and

1-6 m wide. Large pools (0.5-1.8 m deep, 5-6 m long) were

quite common, many with undercut roots, fallen tree limbs

and trapped twigs and leaves lying above or at the water

surface (concealing much of the pool.) Bass Canyon Creek

was the largest of the four streams sampled.

Sonoita Creek originates in the

near the town of Sonoita (T20S, R16E) and flows southwest

to the Santa Cruz River (T22S, R13E) via Patagonia Lake, a

man-made reservoir. The perennial portion of the stream

begins at the town of Patagonia (T22S, R16E) and is composed of groundwater as well as treated municipal

wastewater. These sources provided a mean discharge of

7300 cfs (~2400 cubic meters/second) from 1965-67 (United

States Geological Survey, 1975). The study site was located on the Circle Z Ranch at an elevation of 1350 m and was over 1.1 km long. The upper creek was characterized by long shallow riffles and a number of pools (Appendix A). 12

Pools varied in size and substrate composition, but commonly had fallen tree limbs, twigs, leaves, etc.

(subsequently referred to as debris) at or above the water surface which concealed large portions of the pools.

Streamside vegetation consisted of emergent grasses as well as some small emergent broadleaf plants. The creek below the dam was characterized by slow-moving, murky water, silt/mud covered gravels, lack of pools and reduced riparian vegetation. METHODS 13

Field work began in May 1987 and was completed in

October 1988. One or two trips per month were made except for mid summer 1988 when field trips were curtailed because of heavy precipitation and flooding. Water temperature and dissolved oxygen were measured with a

Yellow Springs Model 54-A oxygen-temperature meter.

Readings for pH were made using a portable VWR digital pH meter. Hydrogen-ion concentration (pH), water and air temperature, and weather conditions were measured on each trip, but dissolved oxygen was analyzed on an intermittent basis. Habitat use was based on fish observed or captured by electroshocking with a Smith-Root Mark VII backpack electroshocker. Most fish were observed and released, but an average of 10 fish per month were collected. Fish collected were placed in a cooler, with chilled water from the stream, and transported to the University of Arizona where they were weighed and measured. Total length was measured to the nearest millimeter using a measuring board and weights were recorded to the nearest gram using an

O'Haus digital scale. Digestive tracts and reproductive organs were removed. Dissected organs and fish (from

Sonoita Creek) were preserved and numbered for cross- reference. Fish from Bass Canyon Creek were frozen and later examined for dietary and reproductive analyses.

Sonoita Creek suckers and dissected organs were 14

preserved in 10% formaldehyde solution for at least 24 hrs

and then transferred to either 70% ethyl alcohol or 70%

Isopropyl alcohol. Sex was confirmed by examination of

gonads and gonad volume was measured using a graduated cylinder for large specimens and a 6 cc syringe for small ones. Fecundity was determined by counting a subsample of

eggs from a known volume of ovary, from 11 fish collected during the spawning season (February through May), with the total number of eggs determined by expanding this value to the total ovary volume.

Stomachs were removed (esophagus to first loop of intestine) from three groups of 10 fish, one group from winter, spring and summer seasons. Contents were identified, where possible, by microscopic examination.

Scales were removed from the left dorsolateral side near the lateral line, anterior to the dorsal fin. Scale imprints were made on plastic slides with a scale press.

These imprints were examined visually using a Model

60-A Bioscope.

On July 15, 1988, 61 fish from Sonoita Creek were collected, weighed, measured and released. These data were used for determining age class structure of the sucker population. In addition, a Length-weight relationship curve was developed by graphing weight (in grams) vs. total length (in millimeters.) RESULTS 15

Preliminary data were taken from four streams but most of the data were obtained from upper Sonoita Creek.

O'Donnell Creek and the Babocomari River had low numbers of fishes and Bass Canyon Creek could not be sampled adequately due to logistical complications.

In Sonoita Creek, water temperature reached a minimum of 9°C on December 14, 1987 and a maximum of 32°C on July

15, 1988. Dissolved oxygen was at or near saturation and pH ranged from 6.5-8.6 during the study period.

A total of 93 suckers was collected, preserved and utilized for reproductive analyses. Stomach analysis of 30 of these fish (10 each from winter, spring, and summer) showed that the dominant food was unidentifiable plant matter (presumably algae), and significant numbers of diatoms were present (10-20% frequency of occurrence).

Stomachs of young-of-year fish contained ~70% diatoms (by number) and unidentifiable plant matter.

A Length-weight relationship curve (Figure 1) from the 61 fish captured and released on July 15, 1988 showed that the suckers ranged from 35-150 mm total length and

1-37 g body weight. Most fish were between 95-125 mm (TL)

(10-22.5 g body weight.)

Scales from 15 of the preserved suckers were examined by Bioscope. No annuli were apparent on the scales but two size groups were evident: Suckers 31-70 mm in length and _ 25 - CT>

20 -

_L ± -L •—•• _i L ± ± J i L ± ± J i L 0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 Total Length (mm)

Figure 1. Length-weight relationship curve for desert suckers. (N=61). 1 7

individuals from 81-150 mm long (Figure 2).

In 1988, the spawning season began in May after a

4-month period of enlarged gonads (late January through

April.) Males exhibited prominent caudal and anal fin tubercles as well as a horizontally striped color pattern

(Figure 3) February through mid April, 1988. Concurrently, gonadal volume was an average of 0.33 ml (range

0.1-1.0 ml) dropping to an average of 0.04 ml (range

0.0-0.08 ml) in late April.

Female gonadal volume averaged 1.28 ml (range

0.1-4.2 ml) during the February-April, 1988 period and dropped to 0.10 ml (range 0.04-0.12 ml) in May. Gravid females yielded an average of 1,140 eggs (range 450-2,772)

1-2 mm in diameter during the reproductive season. Fry were first collected in June, 1988. Fry had the same physical appearance as adult fish.

Almost all (97.3%) suckers were located in pools and rarely found in riffles (2.7%) although pools represented only 3% of the study area. Of the total number of suckers captured, 68% were located in debris pools (those with overhanging limbs and collected twigs, leaves, etc.), 13.5% in pools possessing undercut banks, 11% in backwater pools

(almost all fry), and 4.5% in open, deep (>1 m) pools.

Pools that suckers occupied varied from: 1.2-9.5 m in length, 0.3-2.7 m in width, and 7.6-150 cm in depth. Pool substrate varied as well, from almost 100% sand to 30-40% 20 r-

1 I I I I I I I I" " " I I I 1 I I 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160 Total Length (mm)

Figure 2. Length frequency histogram for desert suckers. (N=61). 00 Figure 3. Desert sucker color phases. (a.) represents normal coloration for males and females. (b.) and (c.) depict two patterns that males exhibited during the breeding season. 20 cobble mixed with sand and gravel. Pools inhabited by adult fish did not have silt or mud substrates, but pools used by fry did have these fine substrates. DISCUSSION 21

Habitat availability is important to the life history

of suckers and is paramount to future survival of the

species.

DIET

Dietary analyses yielded limited information because

stomach contents were not always recognizable. Hauser

(1969) had similar difficulties with mountain suckers

(Catostomus platyrhyncus) in southwestern Montana.

Microscopic examination confirmed that the desert sucker

was primarily herbivorous and fed on diatoms. Comparable

results were obtained by Williams and Williams (1983) for

c. (P.) clarkii in east central Nevada. I observed suckers

feeding off of rocks and gravel, apparently removing

attached algae and associated organisms, in both Bass

Canyon and Sonoita Creeks. Usually numerous suckers fed as

a group in the same area and sometimes on the surface of the same boulder. In Bass Canyon Creek, suckers were generally observed feeding off of larger rocks and

boulders, perhaps because it was easy to scrape diatoms from the large hard surfaces. None of the dissected suckers from Bass Canyon Creek had empty stomachs and only one sucker's stomach from Sonoita Creek was empty. This may indicate that food is not a limiting resource.

Young fish fed heavily on diatoms. Adults probably ate fewer diatoms, thus a dietary shift may occur between 22

fry and adult life stages. However this conclusion is speculative since a large percentage of adult stomach contents was unidentifiable plant matter.

AGE CLASS STRUCTURE

Suckers could not be aged, because annuli were not evident. In many cold-water areas, fish growth stops in winter and an annulus forms. Desert suckers do not experience extreme differences in winter and summer temperatures in southern Arizona, therefore growth is even and scales do not show clear annuli. Greenfield (1970) had similar findings with Santa Ana suckers (Catostomus santaanae) in the Santa Clara River (Los Angeles County)

California. Additionally, the scale method was found to be inadequate by Beamish (1969) for aging white suckers

(Catostomus commersoni) beyond 5 yrs of age at George Lake,

Ontario, Canada. I attempted to identify age groups by length (Figure 2), but there were basically only two groups: young-of-year and fish greater than or equal to

1 yr of age. The older fish ranged from 81-150 mm total length (TL). This group most likely contained a number of age classes.

I was unable to determine the maximum age of desert suckers for Sonoita Creek. Suckers vary in this regard.

Beamish (1969) found a white sucker over 14 yrs of age, while Greenfield (1970) hypothesized that Santa Ana suckers rarely make it past 2 yrs. REPRODUCTION 23

Desert suckers apparently spawn in May and possibly

into June. This time span is similar to that found by

Hauser (1969) for mountain suckers. I never observed

spawning, therefore I was unable to describe what part of

the stream, what type of substrate and what time of day

eggs are laid. Some species of suckers, such as the

longnose sucker (Catostomus catostomus), move from pools to

gravelly, tails of riffles to spawn (Edwards, 1983).

Male and female suckers had enlarged gonads as early

as February 1, 1988, well before spawning took place. I

observed color change, appearance of tubercles on both anal

and caudal fin rays and increased gonad size in males, and

presence of eggs in females (with a concurrent increase in

gonad volume) in February through April. Color changes or

other secondary sexual characteristics were not observed in

females throughout the study period. Combining the above

indicates that spawning took place only once during a

fairly short period of time. Most previous information

concerning this fish's reproductive biology is of a general

nature and largely anecdotal. For example, Minckley (1973)

stated, "Spawning is generally in late winter and early spring, on riffles and in a manner similar to other species of Catostomus." Much remains to be learned regarding this sucker's reproductive habits. HABITAT USE 24

Desert suckers were found almost exclusively in pools with ~80% cover (e.g. visual obstruction from above the water surface) and high water flow in, around and/or through the pools. Maintenance of these pools appears to be one key to their future survival in Sonoita Creek. They seldom occurred in backwaters or riffles, even though some backwaters had a high percentage of cover. Perhaps the missing constituent was sufficient water depth or water flow. Stewart (1926) was unable to find white suckers in pools isolated from inflow and hypothesized that lack of water movement was the reason.

Generally, the pools used by adult suckers had a high percentage of visual cover provided by dead, fallen trees with many branches and trapped debris. This limb/debris layer formed an upper "canopy" that hid fish from view.

Some pools lacking canopy but having undercut banks, large rock piles, and depths of 1.0-1.2 m were also frequented by suckers; however, this type of habitat occurred less frequently than the fallen tree "debris" type of cover.

Similarly, Propst (1982) found that there was a high correlation between amount of pool cover and white sucker population size.

Fry were almost always found in backwaters or water filled depressions, 5 to 10 cm in diameter, adjacent to the main riffle. Perhaps sucker fry cannot maintain position 25 in other parts of the stream due to the strong current and their small size. Harvey (1987) found that young-of-the- year centrarchids and cyprinids could not maintain position in strong currents.

Each pool sampled contained approximately the same number of fish throughout the year, thus my observations gave no indication of any "migration" or seasonal movement.

Bestgen, et al. (Unpubl. data) also found desert suckers to be "sedentary" in the Gila River drainage of New Mexico and

Arizona. Contrary to this, Siebert (1980) found that desert suckers moved away from a gorge in winter, and into the gorge in summer in . He believed that the latter movement was to avoid high water temperatures.

Conflicting results such as these may indicate that there is a high degree of variability in movement patterns of suckers. CONCLUSIONS AND MANAGEMENT IMPLICATIONS 26

Many of the streams previously thought to support

desert suckers contained few fish. Attempts to collect

desert suckers in O'Donnell Creek, the Babocomari River and

the Santa Cruz River produced few or no fish.

O'Donnell Creek is small with narrow riffles, small

pools and relatively slow-moving water. There are few

pools with overhanging logs, branches and other debris.

Desert suckers were present but in very low numbers. There

appears to be little habitat for desert suckers.

In three sampling trips to the Babocomari River I

captured smallmouth bass, Micropterus dolimieui: bluegill,

Lepomis macrochirus; redear sunfish. Lepomis microlophus)

but no suckers. These introduced fish can negatively effect native fish populations by predation (Minckley,

1973). Furthermore, the very restricted size of the river, extremely slow flow rate, silt/mud substrate, indicate that this river may no longer provide suitable habitat for desert suckers.

Bass Canyon Creek contained many desert suckers and the critical habitat features identified in this study.

Upper Sonoita Creek, although not as large as Bass Canyon

Creek, also contained numerous suckers and considerable suitable habitat. Neither of these streams has direct or continuous flow for access to downstream rivers.

Therefore, these groups of suckers currently may be 27

isolated, thereby eliminating the potential for

interbreeding, genetic exchange and recolonization from

downstream. If these isolated habitats are reduced further

or are subject to catastrophic events, elimination of

suckers from these streams could occur.

Protective measures may be required for long-term

preservation of this species and its habitat. Desert suckers have withstood natural changes but have not

adjusted to man-made changes such as the damming of the

Babocomari River or Sonoita Creek. Impoundments may change stream characteristics below dams (Minckley, 1973).

Perhaps slow water flow, settling of fine particulate matter, and changed temperature regimes from impounding these two streams are a threat to both quantity and quality of sucker habitat. Natural changes might allow suckers to adapt to them, but man-induced changes tend to affect natural waters very quickly and radically. The rapidity of these changes preclude adaptation. We need to preserve and enhance suitable habitat to ensure future survival of desert suckers.

One might ask what difference does it make if we lose a species here or there. My answer is, I simply don't know...but perhaps not knowing the consequences is enough.

As Leopold said, "The single most important part of being an intelligent tinkerer, is to save all of the pieces." Appendix A 28

ggnpiUa Creek PqqI Peggription

Pool # Length Width Depth Substrate Type of CQver 1 4m* 3m 1.3m 20%gravel 80%sand QESIl 2 lm 0.3m 0.7m 30%gravel 70%sand undercut bank 3 3.1m 0.7m 0.5m 30%gravel 70%sand undercut bank 4 4m lm 0.5m 20%cobble 10%gravel 70%sand undercut roots 5 8m 1.1 m 0.7m 40%gravel 60%sand debris.fbkwtr) 6 7 m 1 m 1 m 10%cobble 20%gravel 70%sand debris 7 7m 1.5m 0.7m 10%boulder 20%cobble 30%gravel 40%sand debris,rocks 8 11 m 1 m 1 m 40%gravel 60%sand undercut bank 9 4m 2m 0.3m 25%gravel 75%sand debris 10 lm 0.7m 0.2m 40%gravel 60%sand debris 11 4m 0.7m 0.7m 30%gravel 70%sand debris 12 7m 2-2.5 m 0.5 m 10%cobble 20%gravel 70%sand open (bkwtr) 13 12 m 1-2 m 0.3 m 10%boulder 20%cobble 20%gravel 50%sand debris 14 10 m 1-1.1 m 0.25 m 50%gravel 50%sand debris 15 7m 1-1.5m 0.12-0.4m 10%boulder 30%cobble 20%gravel 40%sand debris

* Note that m=meters and bkwtr=backwater. LITERATURE CITED 29

Armantrout, N.B. 1983. Additions to Cross' ecological distribution of the fishes of the . Proceedings of the Desert Fishes Council 9:321-323.

Barber, W.E., and W.L. Minckley. 1966. Fishes of Aravaipa Creek, Graham and Pinal Counties, Arizona. Southwestern Naturalist 11:313-324.

Beamish, R.J., and H.H. Harvey. 1969. Age determination in the 'white sucker. Journal of the Fisheries Research Board of Canada 26:633-638.

Bestgen, K.R., D.A. Hendrickson, D.M. Kubly, and D.L. Propst. Unpubl. data. Movements and growth of fishes in the Gila River drainage, Arizona and New Mexico.

Cross, J.N. 1985. Distribution of fish in the Virgin River, a tributary of the Lower Colorado. Environmental Biology of Fishes 12:13-21.

Deacon, J.C. 1983. The native fishes of the Virgin River. Proceedings of the Desert Fishes Council 8:290-291.

Eddy, S., and J.C. Underhill. 1978. How to know the freshwater fishes. Wm. C. Brown Company Publ. Dubuque, Iowa.

Edwards, E.A. 1983. Habitat suitability index models: longnose sucker. U.S. Dept. of Interior, Fish Wildl. serv., FWS OBS-082/10.8. 21pp.

Fisher, S.G., D.E. Busch, and N.B. Grimm. 1981. Diel feeding chronologies in two Sonoran Desert stream fishes, Agog3,3 chrvsoaaster (Cyprinidae) and Pantosteus clarki (). Southwestern Naturalist 26:31-36

Greenfield, D.W., S.T. Ross, and G.D. Deckert. 1970. Some aspects of the life history of the , Catostomus (Pantosteus) santaanae (Snyder). California Fish and Game 56:166-179.

Harvey, B. C. 1987. Susceptibility of young-of-the-year fishes to downstream displacement by flooding. Trans. Amer. Fish. Soc. 116:851-855.

Hastings, J.R., and R.M. Turner. 1965. The changing mile. University of Arizona Press, Tucson, Arizona. 30

Hauser, W.J. 1969. Life history of the mountain sucker, Catostomus platyrhvnchus. in Montana. Trans. Amer. Fish. SOC. 98:209-215.

Hendrickson D.L. and Simons. 1988. The long hard swim. Wildlife Views, February, 1988:4-7.

Kepner, W.G. 1980. Fishes of the Upper Bill Williams drainage, Mohave and Yavapai Counties, Arizona. Proceedings of the Desert Fishes Council 11:93.

Lowe, C.H., D.S. Hinds, and E.A. Halpern. 1967. Experimental catastrophic selection and tolerances to low oxygen concentration in native Arizona freshwater fishes. Ecology 48:1013-1017.

McNatt, R. 1983. Status of the native Arizona fishes of Aravaipa Creek. Proceedings of the Desert Fishes Council 5:116-117.

Meffe, F.K., and W.L. Minckley. 1987. Persistence of stream community assemblages: pattern vs. causality. Unpubl. Rpt.

Minckley, W.L. 1973. Fishes of Arizona. Arizona Game and Fish Dept. Phoenix, Arizona.

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