Growth, Spawning Preparedness, and Diet of Cycleptus Meridionalis (Catostomidae)

Transactions of the American Fisheries Society 128:900±908, 1999 ᭧ Copyright by the American Fisheries Society 1999

Growth, Spawning Preparedness, and Diet of Cycleptus meridionalis (Catostomidae)

MARK S. PETERSON* Fisheries Ecology Program, Department of Coastal Sciences, Gulf Coast Research Laboratory, Institute of Marine Sciences, The University of Southern Mississippi, 703 East Beach Drive, Ocean Springs, Mississippi 39564, USA

LARRY C. NICHOLSON Gulf Coast Research Laboratory, Institute of Marine Sciences, The University of Southern Mississippi, 703 East Beach Drive, Ocean Springs, Mississippi 39564, USA

DOUGLAS J. SNYDER Mississippi Department of Marine Resources, 1141 Bayview Avenue, Biloxi, Mississippi 39530, USA

GREGORY L. FULLING Gulf Coast Research Laboratory, Institute of Marine Sciences, The University of Southern Mississippi, 703 East Beach Drive, Ocean Springs, Mississippi 39564, USA

Abstract.ÐCycleptus meridionalis is a new species recently distinguished from blue sucker C. elongatus. It occurs in large rivers draining to the northern Gulf of Mexico chie¯y through Alabama and Mississippi. Collections over a 12-month period from the Pearl and Pascagoula rivers, Mis- sissippi, were taken to document the new species' life history characteristics. Female C. meridionalis had signi®cantly greater wet weight (WW, kg) per given total length (TL, mm) (WW ϭ 4.228Ϫ13 TL4.588) than males (WW ϭ 2.000Ϫ9 TL3.237). Aged opercular bones indicated that one annulus formed during September±October at the beginning of the reproductive season. Fish ages in the sample ranged from 6 to 31 years for females and from 4 to 33 years for males. Longevity of C. meridionalis is markedly greater than previous scale aging of C. elongatus had suggested. Male C. meridionalis as small as 327 mm TL (age 4) and females as small as 444 mm TL (age 11) had visibly developed gonads; mean gonadosomatic index was elevated in both sexes between October and March and declined by early April. Spawning began in February to early March when the temperature exceeded 13±14ЊC. The sex ratio was not different from 1:1 (P Ͼ 0.05). Cycleptus meridionalis foraged on aquatic insect larval stages; trichopteran larvae and pupae, chironomid larvae and pupae, and coleopteran larvae were the most common prey items regardless of sex, size, or river. These life history characteristics and food habits are similar to those reported for C. elongatus in the Mississippi River drainage. Our observations indicate that C. meridionalis is viable in both rivers sampled but may be in¯uenced by future river modi®cation as has happened with C. elongatus, which is threatened or endangered over much of its range.

The catostomid genus Cycleptus was long thought a central group in the Mississippi River basin, and to be monotypic, its one species, the blue sucker an eastern group in the Pearl, Pascagoula, and Al- C. elongatus, being limited to large southward- abama river systems. Burr and Mayden (1999) for- draining river systems from Texas to Mobile Bay, mally described the eastern group as a new species, Alabama, and north in the Mississippi River basin Cycleptus meridionalis, for which they suggested to the Missouri and Ohio drainages (Lee et al. ``southeastern blue sucker'' as a common name. 1980). Recently, Burr and Mayden (1999) dis- They left the central group and (for now) the west- cerned three allopatric groups of Cycleptus suckers ern group in C. elongatus. over this range: a western group in the Rio Grande, Cycleptus elongatus was once commercially important in the Mississippi River, but it is now low in abundance and considered threatened or endan- * Corresponding author: [email protected] gered throughout much of its range, due in part to Received June 4, 1998; accepted November 30, 1998 modi®cation of rivers for hydropower and navi- 900 BIOLOGY OF CYCLEPTUS MERIDIONALIS 901 gation (Guillory et al. 1979; Williams et al. 1989; six segments (three in the upper and three in the Tomelleri and Eberle 1990; Elstad and Werdon lower area) for 15 min per segment. 1993). Despite the attention given to its exploi- Two boats and a crew of six were used during tation and subsequent decline, life histories of the collecting. One boat was equipped with a Smith- C. elongatus ``complex'' are only sketchily known. Root electroshocker (model VIA). The voltage and Larval descriptions are available for Mississippi pulses of the electroshocker were set to correspond drainage ®sh (Hogue et al. 1981; Kay et al. 1994). to the conductivity of the water on that date. The Cycleptus elongatus adults exhibit a spawning run shocking boat crew consisted of an operator and upstream in spring, generally in mid to late spring two ®sh dippers. The second ``chase'' boat's crew (Rupprecht and Jahn 1980; Moss et al. 1983; comprised an operator and two ®sh dippers, and McInerny and Held 1988). Moss et al. (1983) ex- the boat was equipped with a live well. The chase amined growth of C. elongatus from the Neosho crew followed the shocking boat, capturing ®sh River, Kansas, based on ages from scales. Once that surfaced behind or alongside the shocking sex was discernible, females were longer and boat. heavier than males. In Pool 20 of the Mississippi All ®sh collected were measured for total length River, Rupprecht and Jahn (1980) examined (TL, Ϯ1 mm), standard length (SL, Ϯ1 mm), and growth using ages based on scales but sexes were total wet weight (TWW, pounds to the nearest 1/ not compared. Food habit studies of C. elongatus 4 pound subsequently converted to kilograms), and revealed that trichopteran larvae and pupae were the presence or absence of breeding tubercles was the most frequent prey, occurring in 97% of all noted. During each electroshocking run, we tab- stomachs in the Mississippi River (Rupprecht and ulated data for all C. meridionalis captured but Jahn 1980) and in 17% in the Neosho River (Moss only kept two or three ®sh per run on any date. et al. 1983). Moss et al. (1983) noted that dipteran From 19 March 1996 to the end of the study, we as well as trichopteran larvae were important di- determined sex of all released ®sh by gently etary components in young C. elongatus. squeezing their abdomen and identifying the ga- Laboratory spawning data, growth rates of mete type. After 2 May 1996 we retained mainly young ®sh (Semmens 1985), and larval descrip- the smallest and largest ®sh to improve our esti- tions are available (Yeager and Semmens 1987) mate of length±weight relationships. Fish that for ®sh from Alabama. However, few published were kept were sexed by gonad examination and data on biology and ecology are available for Cy- their heads and pectoral ®ns were removed, tagged, cleptus meridionalis (Burr and Mayden 1999). To and placed in 95% ethanol. Their body cavities ®ll this data gap, our objectives were to examine were slit open and injected with 100% formalin growth, spawning preparedness, and diet of C. (full-strength formaldehyde), and their bodies meridionalis from the Pascagoula and Pearl rivers, were tagged and placed into 10% formalin. which would provide regional life history data for Laboratory procedures.ÐAfter 1 or 2 weeks, this newly described species. preserved ®sh were ¯ushed of formalin in running tap water. Their entire intestinal tracts and gonads Methods were removed, separated, and blotted, and then General ®eld protocol.ÐWeekly collections of viscera wet weight (VWW, g) and gonad wet C. meridionalis from the Pearl and Pascagoula riv- weight (GWW, g) were recorded. After fat was ers, Mississippi, were taken between late February removed, the viscera were stored in 70% ethanol and early June 1996 (N ϭ 24 collection dates), and until stomach contents could be identi®ed and then monthly until January 1997 (N ϭ 9). Each quanti®ed. Except where noted, all analyses in- river was divided into 36 1.61-km segments,18 in volving wet weight (WW) represent ®eld wet the upper and 18 in the lower general survey area weight (before preservation) minus GWW. Spawn- (Figure 1A, B). For each date, we randomly se- ing preparedness was estimated with the gonado- lected the river order (Pearl or Pascagoula), the somatic index: GSI ϭ 100(GWW/WW) (Crim and general starting area (upper or lower area), and Glebe 1990). All weight values were converted to then the starting segment number (1±18). Once we kilograms prior to analysis. To compare GSI±WW arrived at the starting segment, we electroshocked relationships, we selected only GSIs that were at in a downstream direction for 15 min, dividing the least 1.0% for ®sh collected between October and time shocking between each bank. We collected February, when GSI values were highest. from three consecutive segments; thus on any giv- We used opercular bones to age C. meridionalis en collection date and river, we sampled ®sh from (Scoppettone et al. 1986; Scoppettone 1988). 902 PETERSON ET AL.

FIGURE 1.ÐMaps of (A) the Pearl River, (B) the Pascagoula River in Mississippi, and (C) their regional setting. The 36 1.61-km segments we sampled between February 1996 and January 1997 in each river are numbered 1± 18 above and below the bold lines that separate upper and lower river areas. Boxes with crosses are U.S. Geological Survey gauging stations and arrow heads are launch sites. BIOLOGY OF CYCLEPTUS MERIDIONALIS 903

Opercular bones generally were removed from the order or family and food habits data were pooled left side of the head of each ®sh, but we used the by predator sex and size. right bone if the left one was visibly abraded. After Statistical procedures.ÐWe initially compared the skin of the opercular bone was removed, the several relationships by river and sex with analysis bone was gently cleaned in tap water and placed of covariance (ANCOVA). We could not compare in labeled envelopes for storage. Two independent GSIs between rivers because we did not collect readers examined 187 opercular bones. Measure- ®sh from the Pearl River after August 1996 due ments for marginal increment analysis (MIA, N ϭ to low water nor could we compare age±size re- 161 ®sh) were made with a dissection microscope lationships between rivers because of the inherent along the medial axis of the opercular bone (Scop- variability in age±size relationships for this spe- pettone et al. 1986; Scoppettone 1988) on a line cies. Also, if we separated our age data by river, beginning at the tip of the bony extension dorsal we would not have enough ®sh to validate annular of the opercular±hyomandibular socket to the edge increments. We could, however, compare length± of the operculum. With transmitted light, the weight relationships between rivers. If relation- opaque zone appeared dark and the translucent ships were not different, we used linear regression zone appeared bright (Summerfelt and Hall 1987). on pooled data (river and age) to model relation- Opaque bone is indicative of winter growth, ships between TL and SL, TL and WW, age and whereas broader zones of translucent bone re¯ects TL, age and WW, GWW and WW, and GSI and summer growth. We de®ned an annulus (after WW by sex. The last comparison was used to eval- Scoppettone 1988) as the region in which the outer uate if GSI was a function of WW. If no relation- edge of opaque bone borders the inner region of ship existed, we plotted GSI by sex and date to translucent bone. Percent marginal increment was estimate spawning preparedness. Also, ANCOVA calculated 100(distance from the last formed an- was used to compare male with female slopes and intercepts for the following relationships: TL±WW nulus to the edge of the opercular bone)/(distance (with TL as the covariate), age±TL (with age as between the last two formed annuli) (Crabtree et the covariate), and age±WW (with age as the co- al. 1996). Mean percent marginal increments variate). If the TL±WW relationships delineated (ϮSE, N ϭ 5±61 individuals/month) for all ®shes in the ANCOVA analysis were signi®cantly dif- (rivers and sexes pooled) were plotted against ferent between sexes, their slopes and intercepts month to determine if one annulus (as de®ned) was from the TL±WW were used to estimate their pow- indeed laid down annually and the month in which er functions. If the relationships did not differ, it formed. Narrow increments indicate the male and female data were pooled. The power month(s) when annuli were laid down. There were function is WW ϭ aTLb, for which a is the inter- not enough ®sh per month to consider separating cept and b is the slope of log10-transformed data ®sh into age-classes for the MIA. (presented as back-transformed values) (Anderson One problem of aging catostomids with oper- and Neumann 1996). Chi-square was used to de- cular bones is that growth of support bones in older termine if the compare sex ratio of ®sh captured ®sh hides some of the ®rst-formed annuli (Scop- prior to 2 May 1996 differed from a 1:1 ratio. All pettone et al. 1986; Scoppettone 1988). Because data were log10-transformed when necessary to we collected only two ®sh smaller than 400 mm stabilize homogeneity of variance and normality TL, we borrowed three ®sh smaller than 200 mm assumptions. Relationships were considered sig- TL. Two ®sh (MMNS collection 20130) came from ni®cant if P Ͻ 0.05. the Pascagoula River and one ®sh (USM collection 16731) from the Pearl River (museum codes are Results given by Poss and Collette 1995). We processed During this study, we observed 263 C. meri- the opercular bones from these ®sh and measured dionalis in the Pearl River and 296 in the Pasca- the distance to annuli 1, 2, and 3 along the medial goula River, but we used only the collected indi- axis as noted above. The three values were aver- viduals for various analyses as noted below. For aged for each annulus and used to estimate the females, there was no difference in the TL±(WW presence of hidden annuli in older ®sh. Ϫ VWW) relationship between rivers (ANCOVA: Contents of the stomach (the ®rst descending parallelism, P ϭ 0.366; intercept, P ϭ 0.944), but section of the intestine: Rupprecht and Jahn 1980; for males, this relationship differed (ANCOVA: Moss et al. 1983) were expressed as the frequen- parallelism, P ϭ 0.017; intercept, P Ͻ 0.001). cies of occurrence. Prey items were identi®ed to Mean adjusted male WWs were 14.3 kg for the 904 PETERSON ET AL.

FIGURE 2.ÐMarginal opercular increments (mean Ϯ SE) of Cycleptus meridionalis (pooled by sex, age, and river) by month, January (J) through December (D). There are no data for February; otherwise, means represent 5±61 ®sh per month (N ϭ 161).

Pearl River and 11.2 kg for the Pascagoula River. We do not think this difference, given our ran- domized collection procedures, is biologically meaningful, so we pooled all TL and WW data across rivers for subsequent analyses. The TL±SL FIGURE 3.ÐPlots of body size characteristics in re- relationships were signi®cant for both sexes (fe- lation to age for female (ⅷ) and male (⅜) Cycleptus males: SL ϭϪ56.041 ϩ 0.933TL; r ϭ 0.973; P meridionalis: (A) total length (TL) versus age for 81 Ͻ 0.0001; N ϭ 91; males: SL ϭϪ23.683 ϩ females and 104 males; (B) wet weight (WW) versus 0.876TL; r ϭ 0.962; P Ͻ 0.001; N ϭ 130). Somatic age for 76 females and 107 males. Plotted lines are based WWs of both females and males increased with TL on least-squares regression.

(females: log10TL ϭϪ12.374 ϩ 4.588 ´ log10WW; r ϭ 0.739; N ϭ 90; males: log10TL ϭϪ8.691 ϩ to 593 mm TL (age 33). The relationship of age 3.237 ´ log10WW; r ϭ 0.531; N ϭ 126). However, to TL was positive for both genders (females: TL the slopes of this relationship differed between ϭ 452.246 ϩ 5.535 ´ age; r ϭ 0.638; P Ͻ 0.0001; sexes (ANCOVA; P Ͻ 0.05), the power functions N ϭ 81); males: TL ϭ 450.156 ϩ 4.051 ´ age; r ϭ being WW ϭ 4.236Ϫ13 TL4.588 for females and WW 0.548; P Ͻ 0.0001; N ϭ 104) (Figure 3A). Male ϭ 2.000Ϫ9 TL3.237 for males. and female slopes were parallel, according to The borrowed MMNS ®sh were age 3 and the ANCOVA (P Ͼ 0.05), but there was no signi®cant USM ®sh was age 4. Opercular age agreed with difference (P Ͼ 0.05) in age between males and scale age, and operculum values were used to es- females when adjusted for TL. Thus, growth pat- timate the number of missing annuli on all ®sh. terns did not differ between males and females as One annulus was formed on the opercular bone estimated by age±TL relationships. Relationships during September±October each year (Figure 2). between age and WW also were positive (females:

This indirect method of validation indicates that log10WW ϭϪ0.954 ϩ 0.930 ´ log10age; r ϭ 0. 519; the rings on the opercular bones examined rep- P Ͻ 0.0001; N ϭ 76; males: log10WW ϭϪ0. 416 resented annual rings. The increased marginal in- ϩ 0.457 ´ log10age; r ϭ 0.555; P Ͻ 0.0001; N ϭ crement between June and August indicates sum- 107) (Figure 3B). In this case, slopes differed be- mer to be the primary growing season. tween sexes (ANCOVA; P Ͻ 0.002), the female Females collected in this study ranged between slope being the steeper. 452 mm TL (age 6) and 702 mm TL (age 31), The sex ratio of C. meridionalis did not differ whereas males ranged from 327 mm TL (age 4) from 1:1 (chi square ϭ 2.40; P Ͼ 0.05). There was BIOLOGY OF CYCLEPTUS MERIDIONALIS 905

FIGURE 4.ÐPlots of gonadosomatic indexes (GSI, %) in relation to wet weight (WW, kg) of (A) female and (B) male Cycleptus meridionalis collected between Oc- tober and February when GSI was at least 1.0%. Sym- bols represent months: ⅜ ϭ October; ࡗ ϭ November; ᭡ ϭ December; □ ϭ January; ϫϭFebruary. Plotted lines are based on least-squares regression. FIGURE 5.ÐGonadosomatic indexes (%; mean Ϯ SE) of (A) female and (B) male Cycleptus meridionalis by a strong relationship between GWW and WW be- month, pooled by river and age, and (C) temperature during collections. Means represent 3±26 females and tween October and February, when GSIs were 2±51 males per month. 1.0% or more, for females (r ϭ 0.51; P Ͻ 0.02; N ϭ 24) and males (r ϭ 0.80; P Ͻ 0.001; N ϭ 20). However, there was no relationship between GSI (Ն1.0%) and WW for females (r ϭ 0.077; P TABLE 1.ÐFood habitats (frequencies of occurrence) of Ͼ 0. 05; N ϭ 24) or males (r ϭ 0.032; P Ͼ 0.05; Cycleptus meridionalis (327±702 mm TL; N ϭ 206) from N ϭ 20), indicating GSI was not a function of WW the Pascagoula and Pearl rivers, pooled by length, sex, and date. in either sex (Figure 4). Mean GSIs were elevated for females (up to 20.8%) and males (up to 8.4%) Pascagoula River Pearl River between October and March (Figure 5). They de- Food item (N ϭ 106) (N ϭ 100) clined in early April for both sexes and increased Tricoptera larvae 92 79 again in September (females) and October (males). Tricoptera pupae 23 21 This indicates that spawning occurs in February Chironomid larvae 91 66 Chironomid pupae 7 12 to mid-March, when temperatures increased above Ephemeroptera 42 19 13±14ЊC. Coleoptera adults 37 1 There was no obvious seasonal, gender, or size Empididae larvae 28 8 Plecoptera 25 23 pattern of food consumption or of empty stomachs. Nematoda 24 12 Thus we pooled 206 stomachs across season, sex, Sand 11 22 and TL and expressed data as frequencies of oc- Detritus 10 17 currence (Table 1). Regardless of river, individuals Empty 12 19 906 PETERSON ET AL. mainly consumed trichopteran larvae (83.0%) and 11) had visibly developed gonads. The species pupae (21.3%), chironomid larvae (76.2%) and pu- spawned between February and mid-March, when pae (9.2%), coleopteran larvae (32.5%), and nem- temperatures rose above 13±14ЊC. Although GSI atodes (17.5%). Sand (16.0%) and leaf detritus was not correlated with WW, GWW was so cor- (13.1%) were also frequently encountered in the related, indicating that larger ®sh have larger go- stomachs. nads and presumably larger clutch sizes and higher annual fecundities. Mean GSI was elevated in fe- Discussion males (up to 20.8%) and males (up to 8.4%) be- The life history data for Cycleptus meridionalis tween October and March. Breeding tubercles be- are similar to those reported for C. elongatus. Lon- gan to appear on the head, body, and ®ns of males gevity of C. meridionalis determined with oper- and females in late October but became infrequent cular bones was the greater, however, exceeding by early to mid-May. In Alabama, C. meridionalis 30 years for males and females. Burr and Mayden spawned from mid-March through late April and (1999) noted that R. E. Jenkins (Roanoke College, breeding tubercles were present on both sexes Virginia) used opercular annuli to age a single 460- (Mettee and Shepard 1997). In Alabama, repro- mm TL C. meridionalis collected in the Cahaba duction began when temperatures exceeded 12ЊC River, Alabama, in 1968 and obtained an age of (Semmens 1985) and reproductive condition was 20±21 years. As Burr and Mayden suggested and greatest when temperatures reached 20±21ЊC our data con®rm for C. meridionalis, the longevity (Mettee and Shepard 1997). Northward in the Mis- of both Cycleptus species is considerably greater sissippi River, C. elongatus spawned between late than previous scale aging has indicated. For ex- April to early May at 16.4ЊC in Pool 20 (Rupprecht ample, Rupprecht and Jahn (1980) determined that and Jahn 1980) and from early April at 17ЊC until the oldest scale-based age was 10 (13 from ®n June in Kansas (Moss et al. 1983); hatching oc- rays) for ®sh from the Mississippi River. Moss et curred in early May in Pool 9 of the upper Mis- al. (1983) found female C. elongatus in the Neosho sissippi River (McInerny and Held 1988). As ex- River, Kansas, up to 9 years of age (scale-based; pected, spawning occurs later in northern latitudes. 763 mm TL) and males up to 7 years of age (749 Diets of C. meridionalis did not vary with mm TL). Scales typically underestimate age in ca- length, sex, date, or river and consisted mainly of tostomids and other ®shes (Beamish and Harvey larvae and pupae of aquatic invertebrates. These 1969; Beamish and McFarland 1987). However, prey items are similar to those reported for C. elon- Scoppettone (1988) has validated over an 8-year gatus collected in Pool 20 of the Mississippi River period the use of opercular bones to age cui-ui (Rupprecht and Jahn 1980), in the Neosho River, Chasmistes cujus, an endangered catostomid. Kansas (Moss et al. 1983), and in the Black River, Beamish and McFarland (1987) noted that the use New Mexico (Cowley and Sublette 1987). For ex- of new techniques have increased our understand- ample, Rupprecht and Jahn (1980) documented tri- ing of the longevity of many ®shes, and many chopteran and dipteran larvae and pupae in 97% catostomid and cyprinid species reach ages as of the 46 stomachs examined. Moss et al. (1983) great as 43 years (Scoppettone 1988). Annular determined that adult C. elongatus consumed rings on the opercular bones of C. meridionalis mainly nematodes (66%), trichopteran larvae were laid down between September and October, (34%), and pelecypods (17%), whereas juveniles when water temperatures dropped below 19ЊC and consumed mainly trichopteran larvae (100%) and GSI increased, suggesting reallocation of energy chironomids (100%). and thus mass from growth to reproduction. We only collected two C. meridionalis smaller Female C. meridionalis were heavier per given than 400 mm TL in our study, although we col- length than males. This growth pattern was also lected many small individuals of other bottom- noted by Mettee and Shepard (1997) for C. mer- dwelling species like ¯athead cat®sh Pylodictis oli- idionalis in Alabama and by Moss et al. (1983) for varis, channel cat®sh Ictalurus punctatus, and blue C. elongatus in the Neosho River in Kansas. In cat®sh I. furcatus. These data indicate that small contrast, length±weight relationships did not differ C. meridionalis may occupy a different habitat between male and female C. elongatus from Pool from the main channel and illustrate the need for 20 of the Mississippi River (Rupprecht and Jahn additional research on younger life stages of this 1980). species. Male C. meridionalis as small as 327 mm TL Modi®cation of river systems like the Missis- (age 4) and females as small as 444 mm TL (age sippi River for navigation may have contributed BIOLOGY OF CYCLEPTUS MERIDIONALIS 907 to the present low abundance of C. elongatus by Pages 529±553 in C. B. Schreck and P. B. Moyle, altering spawning runs. This species is considered editors. Methods for ®sh biology. American Fish- threatened or endangered throughout much of its eries Society, Bethesda, Maryland. Elstad, S. A., and S. J. Werdon. 1993. Status report on range. The Pearl and Pascagoula rivers have not blue sucker (Cycleptus elongatus), a candidate en- been modi®ed to the extent imposed on many other dangered or threatened species. U.S. Fish and Wild- medium to large rivers, and the large numbers of life Service, Ecological Services, North Dakota C. meridionalis we collected in each river suggests State Of®ce, Bismarck. viable populations that might be impacted if ad- Gilbert, C. R., Compiler. 1980. Cycleptus elongatus (Le- ditional modi®cation occurs in the future. sueur), blue sucker. Page 396 in D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAl- lister, and R. Stauffer, Jr. Atlas of North American Acknowledgments freshwater ®shes. North Carolina State Museum of This project was funded by the U.S. Fish and Natural History, Raleigh. Guillory, V. C., C. H. Hocutt, and J. R. Stauffer. 1979. Wildlife Service, Section 6 Program, via the Jack- Endangered freshwater ®shes of the southeastern son, Mississippi, of®ce. We especially thank C. United States. Proceedings of the Annual Confer- Knight for the opportunity to conduct this project. ence Southeastern Association of Fish and Wildlife Loans of specimens from the Pascagoula and Pearl Agencies 32(1978):703±714. rivers came from C. Knight (MMNS) and S. T. Hogue, J. J., Jr., J. V. Connor, and V. R. Kranz. 1981. Ross (USM), respectively. We thank J. Christmas, Descriptions and methods for identifying larval blue C. Manning, G. Duff, R. Hendon, G. Crego, C. sucker, Cycleptus elongatus (Lesueur). Rapports et Proces-Verbaux des Reunions, Conseil Permanent Rakocinski, and M. Abney for assistance with ®eld Pour l'Exploration de la Mer 178:585±587. collections. K. VanderKooy assisted with stomach Kay, L. K., R. Wallus, and B. L. Yeager. 1994. Blue content analyses and aging opercular bones. B. sucker Cycleptus elongatus (Lesueur). Pages 114± Jenkins aided us greatly with aging using opercular 121 in Reproductive biology and early life history bones. Nancy Brown-Peterson and three anony- of ®shes in the Ohio River drainage, volume 2: Cat- mous reviewers improved this manuscript. ostomidae. Tennessee Valley Authority, Chattanoo- ga, Tennessee. McInerny, M. C., and J. W. Held. 1988. Collections of References young-of-the-year blue suckers (Cycleptus elonga- Anderson, R. O., and R. M. Neumann. 1996. Length, tus) in Navigation Pool 9 of the upper Mississippi weight, and associated structural indices. Pages River. Transactions of the Wisconsin Academy of 447±481 in B. R. Murphy and D. W. Willis, editors. Sciences, Arts and Letters 76:69±71. Fisheries techniques, 2nd edition. American Fish- Mettee, M. F., and T. E. Shepard. 1997. Status survey eries Society, Bethesda, Maryland. of a blue sucker (Cycleptus elongatus) population Beamish, R. J., and H. H. Harvey. 1969. Age deter- below Millers Ferry lock and dam, Wilcox County, mination in the white sucker. Journal of the Fish- Alabama. Geological Survey of Alabama, Final Re- eries Research Board of Canada 26:633±638. port, Tuscaloosa. Beamish, R. J., and G. A. McFarland. 1987. Current Moss, R. E., F. Harders, and W. H. Tucker. 1983. Ob- trends in age determination methodology. Pages 15± servations on the natural history of the blue sucker 42 in R. C. Summerfelt and G. E. Hall, editors. Age (Cyclepterus elongatus Lesueur) in the Neosho Riv- and growth of ®sh. Iowa State University Press, er. American Midland Naturalist 109:15±22. Ames. Poss, S. G., and B. B. Collette. 1995. Second survey of Burr, B. M., and R. L. Mayden. 1999. A new species ®sh collections in the United States and Canada. of Cycleptus (Cypriniformes: Catostomidae) from Copeia 1995:48±70. Gulf slope drainages of Alabama, Mississippi, and Rupprecht, R. J., and L. A. Jahn. 1980. Biological notes Louisiana, with a review of the distribution, biol- on blue suckers in the Mississippi River. Transac- ogy, and conservation status of the genus. 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