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Home , Creek chubsucker, Golden redhorse, Quillback, Shorthead redhorse, Silver redhorse, Spotted sucker

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111('2,

edf turtle, Chrysemys scripta Differential Use of Habitat by Spawning 4 • :99-108. Catostomidsl er spin Malaclemys terrapin. tic • haviour in free-living KEVIN D. CURRY 2

ra I oostii (Holbrook). Ecol. and 4 . n: M. Harless and H. ANNE SPACIE ,1ey and Sons, . Department of Forestry and Natural Resources, Purdue University, West Lafayette, Indiana 17907 !n n a population of map 'npubl. M.Sc. Thesis, ABSTRACT: Spawning patterns of suckers () in Deer Creek, Indiana, revealed differential use by species and age-size groups. In spring, white suckers c ecology of the map it (Catastornus commersoni) dominated headwaters, while redhorse ( spp.) were i s range. Can. J. Zool., more abundant downstream. Shorthead redhorse (M. macrolepidwurn) were present only in main stem Deer Creek during spawning. Spawning groups of (. f. ans and T. S. Parsons erythrumni) and tM. duquesnet) near the mouth of Deer Creek were rk. significantly older and larger than those upstream in a first-order triburaty. Spawning 11. SAS Institute was not observed until water temperatures stayed above or near 10 C and until water levels were 20-50 cm deep over spawning . Golden redhorse, shorthead redhorse t studies, p. 11-19. In: and northern hog sucker (Hypenteliwn -icans) spawned over medium gravel in riffles wildlife habitat. U.S. 30-50 cm deep with velocities of 0.4-0.9 nilsec. spawned over medium gravel at depths of 20-25 cm where velocity ranged from 0.50-0.59 miser:Creek ehub- (P. suhr4;z). S. Ajr. .1. sucker ( oblongus) select.d fine grave! in areas 50- 75 cm deep having much slower velocity (0.10-0.24 m/sec). (Fos.). 1975. Statisti- rw York. 675 p. orthern population of INTRODUCTION - tributaries serve as spawning and nursery areas for suckers (Catostomidae), ii •435 440. 44 4: • rs:4 in the eastern United •:••but the effect of tributary spawning on maintaining river populations downstream is just beginning to be investigated. Previous reports (Thompson and Hunt, 1930;

.11 lira in northern New Trautman, 1957; Bowman, 1970; Larimore and Smith, 1963; Karr. 197:5; Burr and Morris, 1977; Smith, 1977; and Morrison, 1977) indicate that catostomids in- \1 Graw-Hill Book Co.. habiting ascend smaller for spring spawning. The extent of upstream movements by adults and the degree to which resident and migrant suckers compete tu tles, Genus Graptemys for stream spawning sites are unclear. Lack of specific information on the seasonal use - - V.: Of river tributaries hampers management efforts and an understanding of the potential t black knobbed saw- • populat ions. it Univ., . 81 •c:,' impact of stream alteration on catostomid Studies on catostomid spawning have shown some interspecific differences in the (Al. T.: • My f and Graptemys) in use of river tributaries. Black redhorse (Moxostoma duquesnet) and golden redhorse erythrurum) in the streams and smaller rivers of Missouri and apparently move short distances for spawning (Bowman, 1970; Smith, 1977). Meyer (1962) found no A CEPTED 4 APRIL 1983 evidence for the movement of golden redhorse, shorthead redhorse (M. macrolepidotum) or (M. anisurum) adults into a tributary of the Des Moines River, Iowa, the spawning season. However, the abundance of adult silver redhorse and ( cyprinus) in a tributary of the Hocking River, Ohio, increased dur- 4;..ing spring, suggesting upstream movement into the tributary for spawning (Smith, 04;4 •1977). Some specific measurements of spawning site characteristics for catostomids have f . been reported in recent literature. Black redhorse in Missouri on riffles of rubble •and gravel in 15-60 cm of water (Bowman, 1970). Spotted suckers spawn over course rubble in riffles 0.3-0.5 m deep with flow rates of 1.4 m 3/sec and surface velocities of

. I Journal Paper no. 9585, Purdue Agricultural Experiment Station. 2 Present address: 120 Riverglade, Amherst, MA 01002.

267

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• IV"TRARRI.VMS,ASR,VRT7'. .;*-" • . ,V,41,G9R"14344.4.1,MYS RTR3454=, WMA 268 THE AMERICAN MIDLAND NATURALIST 111(2) x,1984v;f Cc

0.24 m/sec (McSwain and Jennings, 1972). Shorthead redhorse in spawn on ': spring and autumn of 1 similar riffles 15-21 cm deep (Burr and Morris, 1977). Golden redhorse in Clear methods were used to ca Creek, Ohio, spawn over coarse gravel and rubble at velocities of 0.9-1.2 mi sec [-:and a Smith Root Type (Smith, 1977). Whitehurst (1981) has observed spawning activity of creek chubsuckers Smith Root Type VII ba (Erimyzon oblongus) in Duke Swamp Stream, , in 50-80 cm of water. were also collected by ha Other observations of catostomid spawning sites (Pflieger, 1975) suggest that white zones I through III in sc • tr. suckers (Catostomus commersoni), northern hog suckers (Hypentelium nigracans) and creek Presence and nroyem chubsuckers have generally similar requirements and therefore may compete for sites. ing blocknets which were Twelve species of catostomids spawn in Deer Creek, a small tributary of the middle blocknet design of Shette Wabash River in Indiana (Curry, 1979). In this paper we analyze the seasonal seines instead of pipes an distributuion of the most abundant species of suckers in Deer Creek to determine use of sampling period. At eac the creek as a spawning area.-We investigated the distribution of spawning adults, ` 3t upstream, while anotht spawning sequences and spawning site characteristics for the principal species. The downstream. A 2.5-cm ;ta analysis provides new evidence for the differential use of stream spawning areas by n hoopnets and anchored w a.a species and by size-age groups. at zone III because of grt set at noon and checked METHODS Observations were rn: Deer Creek is a third order (Horton, 1945; Strahler, 1957) tributary of the middle tv measurements ++ere ta. 2 Wabash River in Carroll Co., Indiana, which drains 709.7 km of agricultural land. It T. Substrate size was measu 3 has a mean annual discharge of 6.71 m /sec, with daily discharge ranging from adults were identified h+ 0.18-407.6 m3/sec (U.S. Geological Survey, 1977). We sampled six 0.4-0.9 km stream captured in spawning sl sections in four zones (I-IV) on Deer Creek and two of its first order tributaries. backpack shocker. Bachelor Run and Paint Creek (Fig. I..). Zones I and III contained two sample sites We determined the se each. The sampling sites on Paint Creek were smaller in width and had slower velocity or eggs and be the design than those on Bachelor Run or Deer Creek (Table 1). Velocity and discharge were (1957), Pt1iet rr (1975) ar measured directly during the study, using a Pygmy-type flow meter and area - above the bete measurements of cross sections. first two or three fin rays Adult catostomids were collected near the mouth of Deer Creek (zone IV) during .Ages cif >U( krrs were criterion of Spoor (1938). the number of annular ir (Scidn:ore and Glass. 195 of Beatnish (197:3) and Q Comparisons of the at N were made using the Ma; 5~. PAINT CRE K Species 1i,tributinn and mouth of Deer Creek (zt DEER CREEK spawning periods (Table in zone III. seven species captured in zone IV in sin redhorse, and northern he Black redhorse and northt in zones II and Ill. Quill

TARI.F. 1. — Physical char; scale km 1978 Drainage 164.8 8.0 j .usgs area ti Stream (knt'='f o sample Dear Creek 7 09. 7 Bachelor Run 92.9 Fig. 1. — Map of sampling zones 1 through IV on Deer Creek. Bachelor Run and Paint Paint Creek 40.9 Creek, Carroll Co., Indiana. U.S.G.S. = U.S. Geological Survey Gauging Station i

.1 T "1984 CURRY & SPACIE: CATOSTOMID SPAWNING 269 h rse in Illinois spawn on ^spring and autumn of 1977 and 1978 and during summer 1979. Several collecting J iden redhorse in Clear ,`.methods were used to capture adults because of the changes in water level. Hoopnets el cities of 0.9-1.2 m/sec and a Smith Root Type VI boat electrofisher were used during periods of high water. A ti sty of creek chubsuckers Smith Root Type VII backpack shocker was used during lower flow conditions. Adults i. in 50-80 cm of water. ..were also collected by backpack shocker in Paint Creek (zone 1) in spring 1978 and in 975) suggest that white =zones I through III in summer 1978 and 1979. tt sum nigracans) and creek Presence and movements of adult catostomids at zones I-III were investigated us- o may compete for sites. ing blocknets which were set seven times during April-May 1978. We used the general sa 1 tributary of the middle k s blocknet design of Shetter (1938) and Hall (1972) with hoopnets and nylon mesh block e analyze the seasonal - *%:-seines instead of pipes and screening. This allowed blocknet removal at the end of each reek to determine use of sampling period. At each site, one hoopnet set near one bank trapped moving ut on of spawning adults, upstream, while another hoopnet at the opposite bank trapped fish moving th principal species. The downstream. A 2.5-cm mesh blocking seine was stretched diagonally between the stream spawning areas by hoopnets and anchored with reinforcing rods. The stream could not be totally blocked at zone III because of greater depth and water velocity in midstream. Blocknets were set at noon and checked every 4 hr until the morning of the next clay. Observations were made on spawning during spring 1977-1979. Depth and veloci- 57 tributary of the middle ty measurements were taken at each location where suckers were observed spaw ring. k 2 of agricultural land. It Substrate size was measured and classified according to Cummins (1962). Spawning V ischarge ranging from adults were identified by observation through binoculars, by examination of adults pl d six 0.4-0.9 km stream captured in spawning shoals by sport fishermen. and by collection of adults by it first order tributaries, backpack shocker. tained two sample sites We determined the sex and spawning condition of each fish by the presence of milt it and had slower velocity or eggs and b∎ the designated distribution of breeding tubercles specified in Trautman ilo ity and discharge were (1957). Pflieger 11975) and Smith (1977). Scales for age determination were removed tpc flow meter and area above the lateral line between the tip of the pectoral fin and base of they dorsal tin. The first two or three tin rays were also removed from a pectoral tin on each fish. tr reek (zone IV) during a Ages of suckers were determined by annual growth rings on scales following the criterion of Spoor (1938). Ages of selected fish of each species were verified by counting the number of annular marks on thin sections of pectoral tin rays mounted in epos.; (Scidmore and Glass. 1933). Annuli were identified according to the recommendations X of Beatuish (1973) and Quinn and Ross (19831_ Comparisons of the age and length distributions of three species bet en locations were made using the Nlann-Whitnev test (Conover, 1971).

RESULTS CRE K Species. distribution and movements.- More catostomid species were found near the mouth of Deer Creek (zone IV) than in either of its tributaries during the spring spawning periods (Table 2). A total of 10 species were caught in zone IV, nine species in zone III, seven species in zone II and six species in zone I (Table 2). Of 491 adults captured in zone IV in spring, the dominant species were shorthead, golden and black redhorse, and northern hog sucker. These species represented 84% of the total catch. Black redhorse and northern hog sucker were also the most abundant species captured in zones II and III. Quillback were most abundant in catches from zones II and III

TABLE 1.- Physical characteristics of Deer Creek and its tributaries during April and May 1978

Drainage Mean Mean Mean ~~~/ji•usgs area Gradient discharge velocity temperature Stream (km-') (m/ken) ( m './sec) (m/sec) (C) o sample Dear Creek 709.7 1.2 9.6 0.9 13.2 s- Bachelor Run 92.9 2.3 2.4 0.9 13.5 .rk, Bachelor Run and Paint Paint Creek 40.9 1.5 1.1 0.6 12.2 G using Station 5*

1984 270 THE AMERICAN MIDLAND NATURALIST

(Table 2). Almost all of the white suckers were captured in Paint Creek (zone I) and Both golden and black creek chubsuckers were caught only in this zone. Species abundance patterns were very • tured upstream at zon similar between years. Golden redhorsc c The number of species in each zone decreased between spring and summer. younger (P< 0.01) tha Although shorthead redhorse were abundant in spring catches in lower Deer Creek, that the younger, stna they were virtually absent during summer and autumn (Table 2). Relative abundance older, larger adults en of black redhorse in zones I and III also decreased greatly from spring and summer. Northern hog suck White sucker in zone I showed no seasonal change in distribution though their abun- significantly in length. dance increased in summer. Northern hog sucker were abundant throughout the cant. Nevertheless, sp. seasons at all four zones. significantly. smaller (P Blocknet studies at zone II showed that adults moved downstream after dark (Table gests that there is also s 3). The major species were black redhorse, northern hog sucker and quillback. At zone waters do not attract la I, 75% were captured heading downstream, while at zone II, 955 moved Spawning periods and downstream. Blocknet catches at zone III (Table 2) were not as large as those at zones I 1977 than in either 197 and II because the stream could not be totally blocked. Catches did not reflect the upstream and dow•nstn species or abundance of adult suckers we observed in this area during the day. For ex- period and ranged from ample, golden redhorse were uncommon in the blocknet catch at zone III yet w ere crease in stream ternpet abundant on spawning riffles in the same area. Similarly. shorthead redhorse spawned the spring of 1977 and in zone Ill in spring 1978 and 1979 but were never captured at zone III. mained near or above II Size and age distributions. — Size and age distributions were analyzed for the three itiation of spawning ac species found spawning throughout the tributary system: golden redhorse, black Spawning did not begin redhorse and northern hog sucker (Tables 4 and 5). In all cases females were larger in April. when stream tent mean size than males (Table 4). Spawning groups of golden and black redhorse in The combined obser , lower Deer Creek (zone IV) in 1977 and 1978 were significantly larger (P< 0.01) than tivity for suckers in the those captured upstream in zone II (Table 6). This was true when comparisons were sucker, creek chubsuck made on combined data for both sexes and for males and females compared separately. redhorsc, river carpsucke Golden redhorse from zone IV were significantly smaller (P <0.05) in 1978 than in ing in zones I and II. %V 1977. However, black redhorse at the same zone did not differ in size between years. the 1979 spaw ping pent served on spawning area: blocknets TABLE 2. — Percent abundance of all suckers collected in the Deer Creek system he redhOrSe females were in or electrofishing surveys in the spring (Sp), summer (S) or fall (F) of 1977. 1978 or 1979 captured in zone IV dur Zone spawning readiness hut s Changes in l IV species a III reflect the spawning seq. S Sp S F Sp S F Species Sp S Sp redhorse and black redhot black redhorse 32 5 32 8 32 11 8 15 53 0 movement of quillback oc golden redhorse 1 2 8 0 I 24 46 2(1 14 7 93% of this group and In 0 0 0 0 0 0 0 1 0 0 when water temperature shorthead redhorse 0 0 0 0 0 3 3 31 3 0 0 0 1 3 2 4 0 0 silver redhorse 0 0 TABLE 3.—Percent of - 0 0 1 0 4 0 0 - 1 0 0 the spotted sucker piing period for each samplir. white sucker 44 80 1 0 2 0 0 1 0 0 1 % n. hog sucker 21 12 34 73 40 59 33 18 30 65 quillback 3 0 24 19 16 0 8 4 0 0 0. 28 0 0 0 0 4 0 0 6 Date 12-1600 0 0 1 0 0 0 0 0 0 0 creek chubsucker 1 1 0 0 0 0 0 0 0 0 4 April (52) collection years cd cd c d or cd bc bcd d b 14 April Total number (15) captured 283 145 366 74 96 123 118 491 30 29 18 April Number of (31) 3 collections 13 4 8 1 8 3 2 16 4 2 May (88) 1 10 May (8(i) a partial blocknet 18 May (85) b collected in 1977 24 May (7) c collected in 1978 Total (364) <1 d collected in 1979 1984 CURRY & S1'ACIE: CATOSTOMID SPAWNING 271

eek (zone I) and Both golden and black redhorse from zone IV were significantly older than those cap- ured p• tterns were very F upstream at zone II (P<0.01 and P<0.05, respectively). ;: Golden redhorse captured at zone III in the autumn were significantly smaller and i g and summer. " oun er P<0.01 than those taken in the spring at either zone II or IV. This suggests to 'er Deer Creek, ' that the younger, smaller golden redhorse reside in the Deer Creek system while the Le ative abundance older, larger adults enter the lower main stem only for spawning. rri ig and summer. Northern hog sucker spawning groups from zones II, III and IV did not differ th nigh their abun- significantly in length. Age differences between zones II and IV were also nonsignifi- n throughout the cant. Nevertheless, spawning hog suckers in the first order tributary (zone I) were significantly smaller (P <0.01) than those from the other zones downstream. This sug- ai fter dark (Table gests that there is also size segregation among spawning hog suckers and that the head- ! q illback. At zone 3e_ waters do not attract larger adults. • U , 95% moved Spawning periods and species sequence. — Spawning occurred almost 1 month earlier in r • those at zones I 1977 than in either 1978 or 1979 due to an early warming trend. Water temperatures di not reflect the upstream and downstream did not differ by more than 1 C during each spawning ng the day. For ex- period and ranged from 10-22.5 C (Fig. 2). Spawning activity began after a rapid in- z ne III yet were crease in stream temperature above 10 C. Although two such peaks occurred during I r dhorse spawned the spring of 1977 and 1978, spawning did not begin until water temperatures re- on III. mained near or above 10 C for some time (Fig. 2). These temperature increases and in- lur ed for the three itiation of spawning activity followed large decreases in stream discharge (Fig. 3). t.•n redhorse, black Spawning did not begin in 1978 until after the second peak in stream discharge in late tral s were larger in April, when stream temperatures staved near or above 10 C (Figs. 2 and 3). i 1 ack redhorse in The combined observations for :3 years revealed a regular sequence in spawning ac- fixity for Buckets in the following order: white sucker, silver redhorse, northern hog -aT (P<0.01) than t c' mparisons were sucker, creek chubsucker, shorthead redhorse, quillback, black redhorse, golden :tit ' aced separately. redhorse, river carpsut ker ( Carpindes carpio). White suckers were only observed spawn- 15) in 1978 than in ing in zones I and II. We observed silver redhorse spawning in late April only during Biz between years. the 1979 spawning period (Figs. 2 and 3). Although black redhorse were never ob- served on spawning areas, spent females were captured on 20 May 1978 when golden k s stem by blocknets redhorse females were just reaching full spawning readiness. Siale river carpsuckers 7. 978 or 1979 captured in zone IV during mid-March 1977 released milt easily and were close to spawning readiness but spawning was not observed until late April. Changes in species abundance in the blocknet catches at zones I and II in 1978 reflect the spawning sequence (Table 7). Catches of northern hog sucker, golden redhorse and black redhorse increased from 18 April to 10 May. A major downstream 53 0 movement of quillback occurred at night on 18 May at zone II. Females comprised 14 7 93% of this group and most were spent. Increased catches of these species occurred 0 0 when water temperature warmed and stayed above 10 C. Blocknet catches at both 3 0 0 0 TABLE 3. — Percent of the total blocknet catch from Bachelor Run caught during each sam- 0 1 0 piing period for each sampling date in 1978. Total numbers caught in parenthesis. ' = less than 1 0 0 1 %0 1: 30 65 4 0 0 Time period 6 0 28 Date 12-1600 16-2000 20-2400 24-0400 04-0800 0 0 0 0 4 April (52) 2 13 be d b 14 April (15) 2 1 ' 4' 1 30 29 18 April (31) 1 2 4 2 1 4 3 2 May (88) 1 3 13 8 - 10 May (86) 9 14 18 May (85) 2 21 - 24 May (7) 1 1 ' Total (364) <1 <6 42 49 <2

~1( 9' b°[ `i kS:7l +' ►.1~' ~yKl 7s,".ā ~ "~! 'ni7+Fi l.. r7t;.y a ~'•Y1~'i+ii..".-'.^F nL'-Lw.:~ii..d.•_'•r~`"3S'ert.1=•fi<<•.i „Jt''.S'1'w.,`r•°~c~.'=. y,.a 272 THE AMERICAN MIDLAND NATURALIST 1981

zones I and II decreased greatly at the end of May (Table 7), coinciding with the areas. but spaw net decrease in spawning activity. defended small. cle Spawning site characteristics. -Spawning sites were located for live species. (;olden Our observation redhorse, shorthead redhorse and northern hog sucker all used similar sites on riffles site selection by th (Table 8). White suckers spawned in shallower water over medium gravel in the itppur spawning shoals fi: r reaches of the Deer Creek tributaries. Creek chubsuckcrs were titund in di.: s:uuc• temperatures were . TABLE 4. -Total length distribution of suckers captured in the Deer Creek s•: ,ten:: had dropped and ve spring (Sp) and fall (F) The substrate u when tish selected it Length (nun) ing ac t i•. itv, so the c Species • Zone Date Sex Range Mt- an :n silt and sand than th suggesting that the golden redhorse IV Sp-77 M 28 289-450 391.3 itia! choice. The pre F 20 320-479 412.4 39 5 determine whether IV Sp-78 NI 11 321-425 358.6 40 . i F 9 328-447 377.8 41.11 , 44, distrihutio ; II Sp-78 NI 10 290-375 322.b 9.7 to and within the F 17 276-379 332.5 25, within :he main stem Creek (zone I). Its : III F-78 M 13 269-331 296.9 15.7 in autumn, sugc es F 17 289-342 317,t' 13.88 as wui! is solder: re,: black redhorse IV Sp-77 NI 14 304-426 379.4 ;1.3 and a!:tu:nn (Cure, , F 16 362-476 419.1 13.2 st mt P er Creek ye autumn. IV Sp-78 NI 11 34g.- t36 3893' to 7 1 system 1..t- iprit:g sp. to. t F 22 336-470 40j. ti c Tiro' ‘%•ltite su, i round -_>idcnts of Sp-78 NI 31 299-395 31:.4 I' 0 II nbser-. -_tisens on mu F 84 :30:-403 3o1 2 16. 1 mover- • nts clitf-rs e [ Sp-78 NI 15 317-364 338.3 18.1 Grr_;tcr diversit% F 52 315-375 344.3 17.8 silniL:r te, the patter ar northern the Deer hog sucker IV Sp-77 NI 27 256-350 302.:3 25.8 Creek systt F 20 274-382 332.4 .14.5 ( 1963) observed for stickers and creek c III Sp-78 NI 34 250-376 301.4 12.2 2.5.8 km=. Similarly, F 12 280-400 342.3 47.1 catches from zone 1 We found northern f 11 Sp-78 NI 57 235-380 291.1) 34.5 F 28 265-396 334.9 39.1 5. -Age dist! I Sp) and fall (F) I Sp-78 'vI 26 190-332 235.9 32.4 F 12 242-323 273.9 25.5 shorthead Specitn 7. redhorse IV Sp-77 NI 31 364-448 407.7 22.3 goltIcn redhorse F 28 375-479 425.6 29.1 IV - Sp-78 23 370-457 411.8 19.5 F 438.7 21.8 26 396-500 ;ac k redhorse white sucker 24 177-350 240.2 19.9 40 182-409 279.2 55.6 !:. hog ,uc ker quillback II Sp-78 NI 4 305-385 352.3 35.7 F 60 281-444 372.8 39.0

0:3;

U$tALIST 1984 CURRY & SPACIE: C,yrosrowu SI'awasa; 273

(Table 7), coinciding with the = .areas, but spawned in deeper water with slower velocity. Chubsucker males actively defended small, cleared depressions on the sand bottom of pools just above riffles. opted for five species. Golden Our observations suggest that velocity and depth may be key factors in spawning all used similar sites on riffles site selection by these species. During spring 1979, adult redhorse were present at e^ medium gravel in the upper spawning shoals for several days in early May when water levels were high and water ckers were found in the same temperatures were 16-20 C. Spawning did not begin on these shoals until stream levels I in the Deer Creek system during had dropped and velocity decreased. The substrate used for spawning (Table 8) was not the original substrate present when fish selected their spawning areas. All five species clean out areas by their spawn- Length (mm) ing activity, so the gravel uncovered at the spawning sites was larger and more free of Range Mean sn silt and sand than the original material. This was especially true for creek chubsuckers, 1

Creek, Bachelor Run and Paint Creek. supporting Larimore and Smith's (1963) find- Age and size distribute km'. The w T adult ing that northern hog suckers occurred within drainage areas of 6.4 to 823.8 suckers may ere smallest drainage basin areas where Larimore and Smith (1963) collected shorthead 'Y; movements reported in 2. Golden redhorse were most abundant within redhorse redhorse ranged from 205.9-823.8 km reported by Get the range of 205.9-823.8 km' but were also present in small numbers in streams with the younger smaller resit 2 25.8-103.0 km of watershed. Our collections support their observations. Shorthead (1977) did not find app redhorse were observed in Deer Creek 12.8 km upstream from the mouth but never in s populations of golden rc Bachelor Run nor Paint Creek. Golden redhorse were present in the tributaries (zones populations might be pre; I and II) of Deer Creek but were more abundant in the main stem (zones [II and IV). and Spacie (1978) probaE The minimum drainage area of catostomid distribution may represent the approx- on nearby riffles in addi imate limit of stream size required for spawning. Thus, species-drainage area correla- tributaries from rivers. T tions may he useful to assess the amount of potential habitat used by particular species movement upstream. for spawning in river tributaries. Recent work on annul some important questior catostomids. Quinn and F TABLE 6. —Comparison of length and age distributions for spring (Sp) and fall (Fl samples of between age determination adult suckers by location using the Mann-Whitney test. T = Mann-Whitney test statistic. 5 and that fin rays were di Critical values greater than 0.05 were considered nonsignificant (NS). ' = significance at a 0.05; age 7 and older did " = significance at a 0.01 not Carostomids in the Deer Ct - Species Zone Date. N T Significance fish older than age 5 greater a high percentage of other ef Length

golden redhorse II Sp-78 27 106.5 SPAWNING IV Sp-77 53 Ii Sp-78 27 421.5 1979 IV Sp-78 20 1978 IV Sp-78 20 319.0 Iv Sp-77 53 1977 III F-78 30 60.5 Iv Sp-77 53 )24 III F-78 30 573.5 0 •-.,1977 IV Sp-78 20 W20 a--~1978 black redhorse II Sp-78 115 2847 IV Sp-77 30 o--ol 9 79 II Sp-78 115 319.4 ~ 16 IV Sp-78 33 IV Sp-77 30 526.5 NS IV Sp-78 33 Š12 northern hog ~ 8 sucker I Sp-78 38 100.5 ✓ IV Sp-77 47 lr .. W I Sp-78 38 374.0 ā 4 II Sp-78 85 3 II Sp-78 85 2232 NS IV Sp-77 47 0 Age 16 '24 golden redhorse Il Sp-78 23 285 MARCH IV Sp-77 47 Spring water GrePk1durio temperan 51 834.5 g 1977, 1978 and 1979 black redhorse • II .Sp-78 sucker. HS = I\' Sp 77 242 northern hog sucker. ; P chubsucker, Q. quillback, 8 = black ~"-• .sNAVa~

111(2) X1984 CURRY & SPACIE: CATOSTOMID SPAWNING 275

19 3) find- Age and size distribution. —The differential use of tributaries by size-age groups of 2. The 'adult suckers may explain some of' the conflicting observations on spawning I s orthead ā= ovements reported in the past. The limited movement patterns of golden and black ,.;,,. is t within -t`edhorse reportedP bby ~ Gerkingg (1953 ),Bowman ( 197Q )and Smith (1977 ) mamay re represent resent re • ns with the younger smaller residents that move out of pools to spawn on nearby riffles. Smith S orthead — 7:0977) did not find appreciable size differences between upstream and downstream ut ever in populations of golden redhorse, but his tag recaptures suggested that two distinct tri:s (zones populations might he present. Observations by Larimore and Smith (1963) and Curry [I • nd IV). and Spacie (1978) probably include younger upstream resident populations spawning tie • pprox- on nearby riffles in addition to older, larger adults moving into the lower end of ^ea correla- tributaries from rivers. The combined picture would appear to be a mass spawning a!a species movement upstream. Recent work on annulus formation and age determination in white suckers raises some important questions about the age composition of spawning groups of catostomids. Quinn and Ross (1982) have clearly shown that the percent agreement mples of between age determinations using scales and fin rays drops drastically for fish overage est statistic. 5 and that fin rays were difficult to read for fish over age 7. A high percentage of fish cc.ta0.05: : age 7 and older did not form an annulus each year (Quinn and Ross, 1982). ! f Catostomids in the Deer Creek system over age 5 were difficult to age using scales; for --:. fish older than age 5 greater reliability was placed on ag izn iicance es determined from fin rays. If a high percentage of other catostomids fail to form an annulus each year after age 7,

*P- SPAWNING SEQUENCE Y~d CC 1979 WS S HS SH 1978 WS HS S% R R,

1977 WS HS SH C;i C 24 . c.) 0 1977 _ 1978 w20. 0 0 0-01979 16 . w 2 1 2. w I0 8. cr w - 4 - 3ā

0 1 1 ' 1 e 1 ` 1 1 t I r iete1,1'1'1.1YjrI■Ie1 I h I 1 . I~1 8 16 24 I 9 17 25 2 10 18 26 MARCH I APRIL I MAY Fig. 2.—Spring water temperatures and observed spawning sequences for suckers in Deer Creek during 1977, 197$ and 1979. WS= white sucker, S -.silver tedhorse, CS = river - sucker, EIS— northern hog suckcr,.S1-I = shorthead redhorse. C= = golden redhorse, CC - creek chubsucker, Q= quillback , B = black redhorse 276 THE AMERICAN MIDLAND NATURALIST 111(?) {3984

then the mean age of the spawning groups may be older than what we have observed in this study. Spawning sequence and period. —The catostomid spawning sequence we observed in N Deer Creek supports some of the reports of past investigators. Smith (1977) observed the following spawning sequence for suckers in Clear Creek. Ohio: white sucker, silver redhorse, quillback, northern hog sucker, golden redhorse, black redhorse. In Deer 0 Creek. white suckers and silver redhorse were also the first to spawn, but hog suckers preceded the quillbac•ks. Spent female black redhorse were captured in Deer Creek during spring 1978 when golden redhorse began spawning. This indicated that black redhorse spawned earlier than golden redhorse. June (197 7) found that river carpsuckers had an earlier mean peak spawning date than shorthead redhorse on the upper . This was not true in Deer Creek. Only in 1977 were river carpsuckers in spawning condition before shorthead redhorse, but actual spawning still started after shorthead redhorse had spawned. This segregation of catostomid spawning groups by size and time and habitat use would reduce competition for spawning sites and habitat for development of larvae. Hall (1972) hypothesized that adults moved into smaller streams and headwaters which provided more favorable conditions for larval development and growth, but these areas were not unlimited, White sucker. golden redhorse, shorthead redhorse and northern hogsucker have very similar spawning site characteristics (Table 8) and several of these species were observed using the same spawning habitat. We observed golden redhorse

SPAWNING SEQUENCE WS SH 1978 Q B G WS HS SHG HS 56 1977 Q -1977 48 ---1978 2 40 < 0 T u-1 32 tr 124 0 TL) .5.. l 0 16

8

0 1 I I I I T T T . T T T 8 16 24 9 17 25 3 MARCH APRIL

Fig. 3.—Spring discharge and observed spawning sequences for suckers in Deer Greek in • 1977 and 1978. Discharge dam from U.S. Geological Survey-(1977, 1978). WS = white sucker. S = silver redhorse, CS = river carpsucker, HS = northern hog sucker. SH = shorthead redhorse. C = golden redhorse, CC - creek rhubsucker, Q= quillback, 13 = black redhorse I {° - t- % F.T(7n, 7. 1 v. v. `+ rj .y

* l ?t'1*t;tk' ∎10, y t ttl+ifaa,y ~1 :. r , -.- : F~ w.;44!ZnI,T4A ' .: i „... } y x F ;;. ^' t . TABLE 7. —Percent abundance of each species of the total season catch fir each blocknet study on zone Ī (Paint Creek) and zoneYr'(Bachelor Run) during April and May 1978. Total numbers caught for each species in parenthesis. * = less than 1% Zone I Species 4 Apr 14 Apr 18 Apr 2 May 10 May 18 May 24 May

Black rcdhorsc (58) 3 5 5 18 10 2 (;oldrrt redhorse (1) 1 N. hog sucker (52) 1 7 10 7 8 7 White sucker (15) 2 2 4 2 2 1 CURRY & Quillback (4) 1 2 "Total (130) 6 5 15 34 19 12 10

Zone 11 SPACIE: Black redhorse (1 14) 13 2 3 5 7 ° Golden redhorse (43) 1 3 4 3

N. hog sucker (126) I I 5 14 8 :3 1 White sucker (2) . CATOSTOMID Quillback (87) 1 2 19 Total (372) 14 5 4 <_ 9 5 23 21 <_ 26 53 SPAWNING

TABLE 8. —Spawning site characteristics for five species of suckers in Deer Creek'

No. Depth Velocity sites Species (cut) (n,/s) Substrate Description 6 golden redhorse 30-60 0.40-0.90 medium gravel riffle 6 shorthead redhorse 30-60 0.60-0.90 medium gravel riffle 4 northern hog sucker 35-45 0.4(I-0.56 medium gravel riffle 4 white sticker 20.251 0.!")0-0.5!) medium gravel riffle 4 creek chubsurker 50-75 (1.10-0.24 line gravel pool; sand bottom °Size classifications according to Cummins (1962) 278 THE AMERICAN MiDLAND NATURALIST 111(2)

and shorthead redhorse simultaneously on the same spawning . However, the distribution of golden redhorse extends further upstream than shorthead redhorse and provides habitat that is not used by shorthead redhorse. The segregation of black and golden redhorse spawning groups by size acts to distribute the competition for spawn- ing sites over a greater area of stream, since the larger adults do not expend the energy to move to the upstream spawning areas where the smaller and younger adults are more common. White suckers reduce competition for spawning sites with other catostomid species by spawning very early in the season in the upper headwaters. This would allow their larvae to develop earlier than redhorse in habitat away from the main redhorse spawning areas in the Deer Creek main stein. Understanding these spawning patterns will help evaluate the types of habitat needed for the reproduction and maintenance of sucker populations. Stream altera- tions that affect temperature, flow regimes, substrate type or fish movement may lead to changes in catostomid species composition.

Acknowledgments. - We gratefully acknowledge the assistance of Richard Yant throughout this project. James Gammon, George Libey and Harmon Weeks were invaluable in the final development and analysis of the results. The study was funded by the Office of Water Research and Technology (Project A052-IND), U.S. Department of the Interior.

LITERATURE CITED E.A%SISH, R. J. 1973. Determination of age and growth of populations of the white sucker ( Catostomus commerscnt) exhibiting a wide range in size at maturity. J. Fish. Ref. Board .Can., 30:607-616. BowNIAN, M. L. 1970. Life history of the black redhorse, Moxostoma duquesnei (LeSuer). in Missouri. Trans. .4m. Fish. Soc., 99:546-559. 1RR, B. NI. AND M. A. McRRts. 1977 Spawning behavior of the shorthead redhorse, .aloxostoma macrolepidot'm, in Big Rock Creek. Ibid.. 106:80-82. CONOyER. W. J. 1971. Practical non-parametric statistics. John Wiley and Sons. Inc. New York, N.Y. 462 p. CUMMINS, K. W. 1962. An evaluation of some techniques for the collection and analysis of benthic samples with special emphasis on lotic waters. .4m. Midl. Nat., 67.477-504. CcRRV, K. D. 1979. Use of a Wabash River tributary for sucker (Catostomidae) reproduction. Doctoral dissertation. Purdue University, Nest Lafayette. Indiana. 95 p. AND t SPAdIE. 1978. Distribution of stream in Tippecanoe County. Proc. Indiana Acad. Sci., 87:182-187. ER INC. S. D. 1953. Evidence for the concepts of home range and territory in stream fishes. Ecology, 34:347-365. At.t., C. A. S. 1972. Migration and metabolism in a temperate stream . Ibid., 53:585-604. oRowrrz, R. J. 1978. Temporal variability patterns and the distributional patterns of stream fishes. Ecol. Monotr. , 48:307-321. ✓ 1:lORTON, R. E. 1945. Erosional development of streams and their drainage basins; hydrophysical approach to quantitative morphology. Geol. Soc. Am. Bull., 36:275-370. JUNE. F. C. 1977. Reproductive patterns in seventeen species of warm -water fishes in a Missouri River reservoir. Environ. Biol. Fishes, 2:285-296. KARR, J. R. 1975. Stream basin and terrestrial environment factors in fish community structure, p. 47-61. In: J. Morrison (ed.). Environmental impact of land use on water quality: Progress Report-Black Creek Project. EPA-905/9-75-006. LAKE, J. AND J. MORRISON. 1977. Environmental impact of land use on water quality: Final report on the Black Creek Project. EPA-905/9-77-007-B. 28(1 p. LARIMORE, R. W. AND P. W. SMITH. 1963. The fishes of Champaign County, Illinois, as affected by 60 years of stream changes. Ill. Nat. His!. Surf. Bull., 28:299-382. IeSwAtN, L. E. AND R. M. JENNINGS. 1972. Spawning behavior of the spotted sucker Minytrma / melanops (Rafinesque). Trans. Am. Fish. Soc., 101:738-740. EVER, W. H. 1962. Life history of three species of redhorse (Moxostoma) in the Des Moines River, Iowa. ibid., 91:412-419.

,e:,.:,;.. CURRY & SPACIE: CATOSTOMID SPAWNING 279

..• .• GER, W. L. 1975. The fishes of Missouri. Missouri Department of Conservationjefferson • City. 343 p. S. P. AND M. R. Ross. 1982. Annulus formation by white suckers and the reliability of Pectoral fin rays for aging them. North Am. J. Fish. Manage., 2:204-208. ORE, W. J. AND A. W. GLASS. 1953. Use of pectoral fin rays to determine age of the white sucker. Prog. Fish-Cult., 15:114-115. .01r.ER, D. S. 1938. A two-way fish trap for use in studying stream-fish migration. Trans. 3rd N. Am. Wild!. Nat. Resour. conf , p. 331-338. SMITH, C. A. 1977. The biology of three species of Mo.rostoma (Pisces-Catostomidae) in Clear , Creek, Hocking, and Fairfield counties, Ohio, with emphasis on the golden redhorse, .if. erythrurum (Rafinesque). Doctoral dissertation, Ohio State University, Columbus. 158 p. strrx, S. H. 1954. Method of producing plastic impressions of fish scales without using heat. Prog 16:75-78. pooR W. A. 1938. Age and growth of the sucker, Catostomus comrnersoni (Lacepede). in Lake, Vilas County, Wisconsin. Trans. Wis. Acad. Sci. Art.1 Lett., 31:457-505. _.8TRAHLER, A. N. 1957. Quantitative analysis of watershed geomorphology. Trans. Am. Geophys. Union. 38:913-920. TilomPsoN. D. H. AND F. D. HUNT. 1930. The fishes of Champaign County: a study of the distribution and abundance of fishes in small streams. Ill. Nat. Hist. Sure. Bull . 19:1-101. ,TRAUTMAN, M. B. 1957. The fishes of Ohio. Ohio State University Press, Columbus. 683 p. GEOLDGICAL SURVEY. 1977. Water resources data for Indiana water year 1977. U.S. Geological Survey Water-Data Report IN-77-1. 357 p. 1(1 8 Water resources data for Indiana water year 1978. U.S. Geological Survey Water- Data Report IN-78-1. WHITEHURST, D. K. 1981. Seasonal movements of fishes in an eastern North Carolina swamp stream. p. 182-190. In: L. A. Krumholz (ed.j. Warm water streams symposium. S. Div. Am. Fish. Soc., Lawrence, .

...SUBMITTED 29 DECEMBER 1982 ACCEPTED 23 JrNE 1983