Determinants of Nesting Success in the Pumpkinseed ...

Determinants of Nesting Success in the Pumpkinseed (Lepomis gibbosus): A Comparison of Two Populations under Different Risks from Predation Author(s): Stephanie A. Popiel, Alicia Pérez-Fuentetaja, Donald J. McQueen and Nicholas C. Collins Source: Copeia, Vol. 1996, No. 3 (Aug. 1, 1996), pp. 649-656 Published by: American Society of Ichthyologists and Herpetologists (ASIH) Stable URL: http://www.jstor.org/stable/1447529 Accessed: 24-04-2015 19:13 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/1447529?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references.

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Copeia, 1996(3), pp. 649-656

Determinants of Nesting Success in the Pumpkinseed (Lepomisgibbosus):A Comparison of Two Populations under Different Risks from Predation STEPHANIE A. POPIEL, ALICIA PEREZ-FUENTETAJA, DONALD J. MCQUEEN, NICHOLAS C. COLLINS

AND

We compared pumpkinseed nesting success in two lakes having similar size, location, and water chemistry but with different fish communities. Ranger Lake had well-established populations of smallmouth and largemouth bass and few planktivorous fish, whereas Mouse Lake had no piscivores and many planktivorebenthivores capable of consuming pumpkinseed eggs and larvae. Both lakes had well-established pumpkinseed populations, but nesting success was significantly higher in Ranger Lake. The influences of environmental factors (nesting substratum and depth, temperature, solar radiation, precipitation, and wind velocity) on nesting success were investigated, and only wind-induced water turbulence was important. Although wind events were associated with significant amounts of nest destruction, the effects were the same in both lakes. Nest attacks and male nest defense were higher in Mouse Lake, particularly at night. These attacks came from the dense populations of golden shiners found at Mouse Lake and resulted in the loss of many pumpkinseed nests. We conclude that a predatorinduced cascade indirectly influenced nesting success. At Mouse Lake, piscivores were rare, planktivore-benthivore nest predators were abundant, nest-specific behavioral interactions were numerous and nesting success was low. At Ranger Lake, large piscivores were abundant, planktivore-benthivore numbers were low, and nesting success was high.

EST building and offspring rearing in the pumpkinseed (Lepomisgibbosus) are functions performed by the male (Ingram and Odum, 1942; Gross, 1982). The nests, built along littoral areas, are bowl-shaped excavations that the parental male defends against intruders and attackers. It is not clear, however, whether pumpkinseed nesting success (or survival of the offspring to the free-swimming stage) depends on environmental or behavioral factors. Contrary to the bluegill (L. macrochirus)or the longear sunfish (L. megalotis)(Gross and MacMillan, 1981; Dupuis and Keenleyside, 1988), pumpkinseeds do not build colonial nests for protection against predation; therefore, each territorial male has to deal individually with nest predators. Nesting success in pumpkinseed populations would be determined not only by the presence of adequate nesting habitat (Stevenson et al., 1969; Whoriskey and Fitzgerald, 1985) or by weather conditions (Goff, 1985, 1986; Dupuis and Keenleyside, 1988) but also by the behavioral traits of the parental male (Ridgway, 1988). The habitat may place constraints for nest building, such as availability of suitable nest substratum, particle size (Bain and Helfrich, 1983; Nack et al., 1993), or depth of the littoral area. But, to a certain extent, fish can control this

variable by selecting the best suitable nesting habitat (Popiel, 1994). However, fortuitous variables, such as exposure of nests to sudden weather events, might not be predicted by the parental males. Weather can affect nesting success in different ways. Water temperature (depending on day length and degree days) determines timing of reproduction (Burns, 1976; Ridgway et al., 1991), egg development and larval growth (Osenberg et al., 1988; Goodgame and Miranda, 1993) and can indirectly affect male behavior and promote nest desertion (Wrenn, 1984). Other environmental factors, such as heavy rains and storms, produce flooding and promote fungal growth on eggs and egg asphyxiation from siltation (Dupuis and Keenleyside, 1988). However, the wind is probably the most destructive side effect of weather instability on the nests. Goff(1986) found that, in general, the greater the number of hours a nest was exposed to the effect of wind, the smaller the number of free-swimming larvae produced. From the pumpkinseed perspective, nest predation is probably a more real threat for their offspring. It is in this area where parental care and male brood-defense behavior can minimize the impact of fish predation on the offspring (Coleman et al., 1985; Ridgway, 1988). In fact,

? 1996 by the American Society of Ichthyologists and Herpetologists


COPEIA, 1996, NO. 3

650

ANDRANGER FORTHEMAJOR FISHSPECIES ESTIMATES FOUNDIN MOUSE TABLE1. BIOMASS ANDDENSITY LAKES. Estimatesare based on sequential mark-and-recaptureexperiments. NE = not enough recapturesto make an estimate, NP = not present. All fish were age 1+ or older. Ranger L. Biomass (kg ha-')

Lepomisgibbosus

pumpkinseed

Mouse L.

Density (numbers ha-')

Biomass (kg ha-')

Density (numbers ha-')

4.66

166

10.79

1031

18.70

58

16.17

16

0.23

3

7.08

479

NE

0.70

175

NP

NP

0.06

114

4.37

50

NP

NP

1.51

24

NP

NP

NP

NP

2.60

Catostomuscommersoni

white sucker Perca flavescens

yellow perch Notemigonus crysoleucas

golden shiner

< 0.1

Semotilus atromaculatus

creek chub Micropterussalmoides

largemouth bass Micropterusdolomieui

smallmouthbass Ameiurus nebulosus

brown bullhead

active male nest defense has even been observed at night (Hinch and Collins, 1991), and total nest failure has resulted when guarding males (Micropterus)were removed from the nests (Kramer and Smith, 1962; Neves, 1975). In lakes where fish-fish interactions are high, the success of a nest may depend on how well the male can defend his young and the quality and quantity of his opponents. Comparison of populations under different predation pressures in similar, closely located systems, yields the opportunity to isolate behavioral and environmental factors from general weather conditions. We have tried this approach in the present study. We have simultaneously monitored two reproducing pumpkinseed populations with different rates of reproductive success. Our hypotheses were that the lake-specific patterns in nesting success might be explained by differences in habitat availability for nesting, effects of sudden weather changes on the nests and different levels of shoreline exposure to the wind, and lake-specific patterns of male brood-defense behavior attributable to differences in nest predator abundances. METHODS

Mouse Lake (451 1'N, 78?51'W) and Ranger Lake (40'9'N, 78?51'W) are small, single-basin, oligotrophic, and moderately humic lakes located in south-central Ontario (Sherbourne

27

Township), Canada. The two lakes are morphometrically and chemically similar (Ramcharan et al., 1995). Ranger Lake has a surface area of 11.25 ha and a mean depth of 5.62 m. Mouse Lake is slightly smaller and shallower, with a surface area of 8.99 ha and a mean depth of 4.88 m. Littoral substrata in both lakes are a mixture of sand and cobble with varying amounts of organic material. Mouse Lake has more organic substratum and a more extensive macrophyte cover than does Ranger Lake. The fish communities in these lakes were very different (Table 1). We estimated fish populations using sequential mark-recapture experiments (Schnabel method) for 10 dates both in 1992 and 1993. Ranger Lake had supported large populations of piscivores (largemouth bass and smallmouth bass) for about 20 years (Ramcharan et al., 1995); and golden shiners and yellow perch were rare, numbering only a few individuals ha-'. In contrast, Mouse Lake had remained essentially piscivore-free for 17 years (Ramcharan et al., 1995), and the planktivorebenthivore fish population numbered more than 1200 individuals ha-'. The total biomass of pumpkinseeds was twice as high in Mouse Lake, and the total densities were seven times higher. The Mouse Lake pumpkinseeds weighed less at any given length and reached lower maximum total lengths than Ranger Lake pumpkinseeds (approximately 16 cm vs 26 cm, respectively; Popiel, 1994). The majority of the zooplankton species pres-


POPIEL ET AL.-PUMPKINSEED ent in these lakes were common to both, but zooplankton densities were higher in Mouse Lake (Ramcharan et al., 1995). The benthic invertebrate communities also had similar species compositions, but the Ranger Lake benthic biomasses were higher (Ramcharan et al., 1995). Nesting and reproductionsurvey.-In 1992 and 1993, locations of individual nests and times of establishment were determined from snorkeling surveys (Fig. 1). The surveys started at the beginning of the nesting season when active nests were sighted. Surveys were conducted at fiveto seven-day intervals in 1992 and at two- to four-day intervals in 1993. All nests were numbered and nesting maps continuously updated until the end of the nesting season. On each survey, the condition of each nest was categorized as clear and active, with the male actively maintaining the nest; reproductively active with early stage eggs present (clear or orange in color); reproductively active with later stage eggs present (opaque or white in color); reproductively active with early stage fry present (clear fry); reproductively active with late stage fry present (large, black fry); or inactive, with no male, eggs, or fry present. Three measures were used to quantify nesting success: daily percent successful, daily percent starts, and daily percent failures. Daily percent successful was calculated by dividing the total number of nests with "successful fry" on any given day by the total number of active nests on that same day. Successful fry were those in nests close to stage 5 (black fry) in a survey but not found in the next survey (we assumed they had moved away from the nest between surveys) and those observed in stage 5 (black fry) during the survey. Daily percent starts was defined as the total number of nests started since the last survey, divided by the total number of active nests (all stages) observed on the survey day. Daily percent failure was calculated as the total number of failed nests since the last survey, divided by the total number of active nests observed on the survey day. Failed nests were identified as having gone from an active status on the day of a survey to being abandoned and filled with silt or sediment on the day of the next survey. Physical characteristics.-For both lakes, maps of littoral substrata were created incorporating information from complete shoreline surveys (from a boat and snorkeling). In 1992, nest substratum type and nest depth (distance from the surface of the water to the center of the nest) were recorded by snorkeling. We used the fol-

651

NESTING SUCCESS Mouse Lake June 13, 1993 N

A

0

100 200m

RangerLake June 12, 1993

I

Fig. 1. Maps of Mouse and Ranger lakes. Small circles in littoral area are pumpkinseed nests on 13 June (midreproductionseason).Lakesare dividedinto the eight sectionsused for determinationof wind impact on the nests. Sections were numbered daily depending on cumulative wind direction from 1 (the most downwind)to 8 (the most upwind).


652

COPEIA, 1996, NO. 3

lowing categories (modified from Bain and Helfrich, 1983) to describe both the available nesting substrata in the area where a nest was found and the nest materials where the eggs were laid: fine, medium, and coarse (1) sand-including sand (< 2 mm); (2) mixed-a combination of sand, gravel (2-32 mm), and pebble (32-64 mm) > 64 mm in substrata; (3) large cobble-stones tree branches diameter; (4) sticks-including and twigs; (5) organic-usually a combination of leaves, organic silt, and small branches; (6) boulders-large rocks or part of the cliff areas found in both Mouse and Ranger lakes. Meteorological observations.-Hourly incoming solar radiation (daylight hours), air temperature, wind speed, and wind direction data were obtained from the Ontario Ministry of the Environment and Energy databases (Dorset Centre). All variables were measured hourly at two fixed stations at similar distances from the two lakes. Values from the two stations were averaged together for the 24-h period prior to any given survey. To assess the effect of wind, the lakes were divided into eight sections (45? angles) along the major axes of the lakes (Fig. 1). Predominant winds were north and west, but because the lakes had different orientations (Fig. 1) and topographic features, Ranger Lake was more affected by north winds and Mouse Lake was more affected by west winds. The resultant effect of the wind action on the lakes (i.e., turbulence effects) was highest on the southeastern shore of both lakes. To measure the wind effect, hourly mean wind velocities were associated with each 45? lake section (Fig. 1). Through each day, the hourly average speeds associated with each of the eight 45? lake sections were added. At the end of each day, this process resulted in the estimation of a cumulative daily total wind speed (based on hourly data) for each lake section. These were then compared with rates of nest destruction in each of the eight sections of each lake. Behavioral observations.-In 1993, direct observations of male nesting behavior were made with a time-lapse video recording system (Collins, 1989; Hinch and Collins, 1991). Pumpkinseed nests (bare, sandy circles 30-50 cm in diameter) in Mouse and Ranger lakes were chosen at random and recorded on the same dates for a period of 24 h. The camera was positioned 0.7 m vertically from the bottom and 0.5 m horizontally from a given nest. During the night, video recordings were illuminated using infrared light (quartz-halogen bulbs, Schott RG780 filters) not visible to the fish (Collins and Hinch, 1993).

Images were recorded every 0.2 sec, and playback speed was chosen at 60 fields per second. To quantify different components of nest guarding and defense both night and day, four time periods-night (0000-0200), dawn (06000800), day (1200-1400), and evening (1800viewed from recordings of seven 2000)-were nests from each lake. Four behavioral components were measured for every male monitored: (1) total time spent on the nest-the amount of time the male guarded the nest; (2) mean exit time-the average amount of time spent away from the nest per departure; (3) number of intruders entering within the perimeter of the nest; and (4) number of attacks-number of direct confrontations between the guarding male and individual intruders. Data from observations were ranked, because logarithmic transformations could not correct problems of unequal variances and nonnormal distributions, and analyzed with a twoway parametric repeated-measures ANOVA with lake and time of day (repeated) as the main variables (Zar, 1984; Systat for Windows ver. 5.03, Eceston, IL, 1992, unpubl.). For each variable, all times and both lakes were combined before ranking. Ranks were, therefore, across lakes and times. RESULTS

The summer of 1992 was colder (1992: x = 14.3 C, 1993: x = 17.1 C), had lower total solar radiation (1706.86 kJ/m2/day mean difference), and higher rainfall (1992: 4.86 mm, 1993: 2.88 mm) than the summer of 1993. Average wind velocity was 4.68 km h-' in both years. More total nests were established (not necessarily successful) in 1993 than in 1992, and more nests produced fry in 1993 in both lakes. Percent nesting success with respect to time was consistently higher in Ranger Lake than in Mouse Lake in both years (Fig. 2). Explanations for these trends were sought from the biotic and abiotic factors. The amount of area < 1 m deep in the littoral zone was the same for both lakes (Mouse: 0.745 ha; Ranger: 0.794 ha) and offered comparable substrata for nesting. Most types of substrata were evenly represented in Mouse Lake, whereas Ranger Lake was dominated by mixed substrata (sand-gravel-pebble) and sand (Fig. 3). Pumpkinseed use of habitat for nesting was similar in both lakes (Wilcoxon sign-rank; P = 0.53), and the most selected substrata were mixed, sticks and sand (Fig. 3). Nest depth did not affect nest success in either lake. Frequency distributions of nest depths showed no significant differences between the


POPIEL ET AL.-PUMPKINSEED 100 !90 -

/

10 -

/

5

0

170

160

180

190

200

210

90 -

80 = 70c) 6050-

o 40 3020 10 o 150 30-May

a35 - E B 30 5 25 -

,

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1I

150 100

^ ILl a

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653

NESTING SUCCESS

180 29-June DAYof YEAR

% 20 x15 't O105< 0 C 16(0

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I

I

170

175

165

170

0

lakes (Komolgorov-Smirnov, P > 0.05). Mean depths of successful and unsuccessful nests in Ranger and Mouse lakes were similar (30-40 cm) (Komolgorov-Smirnov, P > 0.05). Nest starts and air temperature were not correlated (Mouse r2 = 0.02, Ranger r2 = 0.38), nor were nest starts and increased solar radiation (Mouse r2 = 0.02, Ranger r2 = 0.14; Fig. 4). Nest destruction was related to wind velocity in both lakes (Fig. 5; Mouse r2 = 0.48, Ranger

160 9-Jun

-

190

175 180 24-Jun DAYof YEAR

185

190 9-Jul

MouseLake'60 '40 e '0 U-

^20

of slopes, t = 1.21; df

1; P > 0.05). Nest failure occurred most frequently at the downwind ends of the lakes and was associated with turbulent destruction. Behavioral variation among nesting males at a given time and place was high so that statistical

~

I

185

Fig. 4. Mean (A) precipitation(bars)and temperature (line) and (B) solar radiation measured by the Ministryof the Environmentand Energyat the monitoring stationsat Paint and Plasticlakes and (C) nest starts (%).Data averaged from 1992 and 1993.

-

[-- 40

I

180

210 29-July

Fig. 2. Pumpkinseednest success rates over time in Mouse and Ranger lakes during 1992 and 1993.

r2 = 0.56; Homogeneity

i

165

'

0 -

Selected qAvailable

Z

Mo_

1'00

WindDirection

10 Cumulative WindSpeed (kmh )

MOUSE

~' 30-

0 ._

z

ULL

0 Mixed

Sticks

Sand

Large cobble

Organic Boulders

t5 z la

30'

i

20-

Mixed

Sticks

Sand

Lare

Organic Boulders

SUBSTRATUM TYPES

Fig. 3. Distributionof substrataavailableand selected for nesting in Mouse and Ranger lakes, 1992.

00

Cumulative 81 WindSpeed (kmh ) Fig. 5. Nest failure plotted as a function of wind direction and log (x + 1) cumulativewind speed for Mouseand Rangerlakes.Peaksin the surfaceindicate highest failure rates; flat or low areas indicate low nest failure rates. Direction 1 is the most downwind; direction 8 is the most upwind. WindDirection6


654

COPEIA, 1996, NO. 3

80

2.

TABLE

PROBABILITY-VALUES

(REPEATED-MEAOF RANKS) FOR THE DIFFERENCES BETWEEN LAKES AND TIMES OF DAY FOR EACH OF FOUR

A

C'

SURES NEST

ci

ANALYSIS

DEFENSE ACTIVITIES.

Asterisks indicate proba-

bilities that are significantafter sequentialBonferroni correction to keep a - 0.05.

o --

E

Source of variation

0

Time on nest

1 0.4 Lake Time of 3 0.0001* Day Lake x Time 3 0.1

?l50 0 cn

0)

o

0

C)

df

Time per Number of absence intruders

Number of attacks

0.06

0.001*

0.0001*

0.02

0.02

0.5

0.8

0.04

0.2

o 25

.0

o C:

0 5 O

300 .,0

co

-1-

150 ..

L

1O -0

.

L-

.

) a)

o

50 Z)

E z

0

R

M R

M

R

M

00001800060012002000 0200 0800 1400 in activities defense four nest Diel variation 6. Fig. by malesout of seven nests filmedfrom each of Mouse and Rangerlakesduring 1993. Squaresrepresentthe medians,verticallines the ranges. In each 2-hourtime block, the data on the left are for Ranger Lake (R) and the data on the right are for Mouse Lake (M).

analysis of ranks; Table 2). Only for "time on the nest" was the time-of-day effect significant (P = 0.0001) after Bonferroni correction for four simultaneous analyses. This effect arises from a relatively higher portion of time spent on the nest at night in both lakes. The time-ofday effect was nearly significant (P = 0.02, critical P = 0.016 with Bonferroni correction). In both lakes, there was more time spent on the nest at night (0000-0200) than at any other time (Fig. 6A). Also, males at Ranger Lake tended to exit the nest for longer periods of time (P = 0.06) during all time periods than the males at Mouse Lake (Fig. 6A-B). The two lakes differed in the "number of intruders" (P = 0.001), especially at night (P = 0.02). Mouse Lake had the highest intruder level (Fig. 6C). The number of attacks on nests was significantly higher (P = 0.0001) for Mouse Lake (Table 2), and the median exceeded that for Ranger Lake in all four time periods (Fig. 6D). Underwater video records revealed that golden shiners were responsible for over 95% of the intrusions, and golden shiners and brown bullheads were responsible for almost all of the attacks. Overall, pumpkinseed males in Mouse Lake tended to spend less time away from the nest per departure and faced far more intruders and more attackers than their counterparts at Ranger Lake. Six of the seven nests observed in Mouse Lake were abandoned by the male sometime during the 24 h of filming. In Ranger Lake, only one of the seven nests was abandoned. DISCUSSION

significance was achieved only for large effects (Fig. 6). There were no significant interactions between lake and time of day for any of the four nesting activity variables (repeated-measures

Although storms and other atmospheric events can be a source of recruitment failure (Eipper, 1975; Goff, 1986), most offspring are usually lost to predators (Bain and Helfrich, 1983; Keenleyside and Mackereth, 1992). In


POPIEL ET AL.-PUMPKINSEED fact, the differences observed in pumpkinseed nesting success in our two study lakes were the result of biotic factors involving different fish densities and fish community structure. Mouse Lake parental males faced a more complex set of fish interactions, involving intraspecific competition and interspecific predation on their offspring, than the males at Ranger Lake. In Mouse Lake, we expected pumpkinseeds to compete for suitable nesting grounds, since population densities were high. We found that 15% of the nests in this lake were built on "less preferred" substrata, such as organic matter and boulders. Although centrarchid males show preferences for certain species-specific depths for nesting (Pflieger, 1966; Gross and Nowell, 1980), the depth range used by nesting pumpkinseed males was broader in Mouse Lake (20110 cm) than in Ranger Lake (20-80 cm). Nest starts were also lower in Mouse Lake than in Ranger Lake, despite a higher density of reproducers. All these observations support the hypothesis that pumpkinseeds at Mouse Lake were under strong intraspecific competition for nesting grounds. Male investment in brood protection was also higher in Mouse Lake than in Ranger Lake because of a higher nest predation pressure. Our results showed that most attacks on the nests took place at night, which agrees with Hinch and Collins (1991), and also showed that males in Mouse Lake had to fend off more intruders and attackers and were forced to remain alert for more hours than their counterparts in Ranger Lake. Coleman et al. (1985) found that parental investment in bluegill is determined by the amount of investment made by the male in a nest and by the actual brood size. In Mouse Lake, the investment in defense was very high, but the brood size decreased as the attacks on the nest increased. We observed that most of the videotaped nests were deserted in Mouse Lake, presumably because the brood size reached some lower limit that unbalanced male investment criteria. In general, abiotic factors had various influences on patterns of pumpkinseed nesting initiation and survival; but in all cases, both lakes were equally affected, and between-lake differences in nesting behaviour were not explained. The most significant climatic variable influencing nesting success was wind-generated turbulence. These results are in accordance with observations by Goff (1986) for smallmouth bass. The only temperature-related difference found between both lakes was a higher lake-specific warming rate in Ranger Lake during earlyspring. Apparently, the warmer water in Rang-

NESTING SUCCESS

655

er Lake was associated with earlier pumpkinseed nest initiation (e.g., Shuter et al., 1980) compared to Mouse Lake. However, late reproduction in Mouse Lake also could have been associated with differential energy reserves characteristic of stunted pumpkinseed populations (Danylchuck and Fox, 1994). Overall, we conclude that, for both Mouse and Ranger lakes, habitat characteristics were similar and weather effects equal. Therefore, between-lake differences in nesting success were primarily determined by piscivore mediated fish community structure which strongly influenced the ability of pumpkinseed males to successfully defend their offspring through to the freeswimming stage. Contrary to our initial expectations, the high densities of piscivores found at Ranger Lake were associated with high rates of nesting success involving rather low rates of parental investment. On the other hand, Mouse Lake males frequently failed in their attempts to produce free-swimming larvae, despite the fact that parental investment was high. Failure did not result from inferior parental behavior but rather because the Mouse Lake males found themselves in a complex, highly competitive fish community comprising large numbers of nest predators. At Ranger Lake, small-bodied nest predators were controlled by piscivores (smallmouth and largemouth bass), and pumpkinseed nesting success was high. ACKNOWLEDGMENTS

We are grateful to K. Hughes for providing statistical analyses and to C. Ramcharan for providing assistance with figures. Many thanks to A. Lahti for assistance in the field and to everyone who works on the Dorset Lakes Research Project. This work was funded by operating and equipment grants from the Natural Sciences and Engineering Research Council (Canada) to DJMcQ and NCC. LITERATURE CITED BAIN,M. B., ANDL. A. HELFRICH.1983. Role of male

parental care in survivalof larvalbluegills. Trans. Am. Fish. Soc. 112:47-52. BURNS, J. R. 1976. The reproductive cycle and its environmentalcontrol in the pumpkinseed,Lepomis gibbosus(Pisces:Centrarchidae).Copeia 1976:449455. COLEMAN R. M., M. R. GROSS,AND R. C. SARGENT.

1985. Parental investment rules: a test in bluegill sunfish. Behav. Ecol. Sociobiol. 18:9-66. COLLINS, N. C. 1989. Daytime exposure to fish pre-

dation for littoral benthic organismsin unproductive lakes. Can.J. Fish. Aquat. Sci. 46:11-15.


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, AND S. G. HINCH. 1993. Diel and seasonal variation in foraging activities of pumpkinseeds in an Ontario pond. Trans. Am. Fish. Soc. 122:357365. DANYLCHUCK, A.J., AND M. G. Fox. 1994. Seasonal

reproductivepatternsof pumpkinseed(Lepomis gib-

bosus) populations with varying body size characteristics. Can. J. Fish. Aquat. Sci. 51:490-500. DUPUISH. M. C., ANDM. H. A. KEENLEYSIDE. 1988. Reproductive success of nesting male longear sunfish (Lepomismegalotispeltastes). I. Factors influencing spawning success. Behav. Ecol. Sociobiol. 23: 109-116. EIPPER, A. W. 1975. Environmental influences in the mortality of bass embryos and larvae, p. 295-305. In: Black bass biology and management. R. H. Stroud and H. Clepper (eds.). Sport Fishing Institute, Washington, DC. GOFF,G. P. 1985. Environmental influences on annual variation in nest success of smallmouth bass, Micropterusdolomieui,in Long Point Bay, Lake Erie. Environ. Biol. Fishes 14:303-307. . 1986. Reproductive success of male smallmouth bass in Long Point Bay, Lake Erie. Trans. Am. Fish. Soc. 115:415-423. L. S., AND L. E. MIRANDA.1993. Early GOODGAME, growth and survival of age-0 largemouth bass in relation to parental size and swim-up time. Ibod. 122:131-138. GROSS,M. R. 1982. Sneakers, satellites and parentals; polymorphic mating strategies in North American sunfishes. Z. Tierpsychol. 60:1-26. ,AND A. M. MCMILLAN.1981. Predation and the evolution of colonial nesting in bluegill sunfish (Lepomismacrochirus).Behav. Ecol. Sociobiol. 8:163174. ,AND W. A. NOWELL.1980. The reproductive biology of rock bass, Ambloplitesrupestris (Centrarchidae) in Lake Opinicon, Ontario. Copeia 1980: 482-494. HINCH,S. G., ANDN. C. COLLINS.1991. Importance

of diurnaland nocturnalnest defense in the energy budget of male smallmouthbass:insights from direct video observations.Trans. Am. Fish. Soc. 120:

657-663. INGRAM,W. M., AND E. P. ODUM. 1942. Nests and behavior of Lepomisgibbosus (Linnaeus) in Lincoln

Pond, Rensselaerville,New York. Am. Mid. Nat. 26:182-191.

M. H. A., ANDR. H. MACKERETH. 1992. KEENLESIDE, Effects of loss of male parent on brood survival in a biparental cichlid fish. Environ. Biol. Fishes 34: 207-212. KRAMER,R. H., AND L. L. SMITH. 1962. Formation

of year classesin largemouthbass.Trans. Am. Fish. Soc. 91:29-41. NACK, S. B., D. BUNNELL, D. M. GREEN, AND J. L. FORNEY. 1993. Spawningand nursery habitats of largemouth bass in the tidal Hudson River. Ibid. 122:208-216.

NEVES,

R. J. 1975. Factors affecting fry production

of smallmouth bass (Micropterusdolomieui) in South

Branch Lake, Maine.Ibid. 104:83-87.

C. W., E. E. WERNER,G. G. MITTELBACH, OSENBERG,

ANDD.J. HALL.1988. Growthpatternsin bluegill

(Lepomismacrochirus)and pumpkinseed (L. gibbosus)

sunfish: environmental variation and the importanceof ontogeneticniche shifts.Can.J.Fish.Aquat. Sci. 43:17-26.

W. L. 1966. Reproduction of the smallPFLIEGER, mouth bass (Micropterusdolomieui) in a small Ozark stream. Am. Mid. Nat. 76:410-418. POPIEL, S. A. 1994. An analysis of the factors regulating growth and recruitment in pumpkinseed (Lepomisgibbosus)sunfish transport model. Unpubl. master's thesis. York Univ. Toronto, ON, Canada. C. W., D. J. MCQUEEN,E. DEMERS,S. RAMCHARAN, A. POPIEL,A. M. ROCCHI,N. D. YAN, A. WONG,

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(SAP, DJMcQ, AP-F) DEPARTMENT OF BIOLOGY, YORK

UNIVERSITY,

4700

KEELE

STREET,

TORONTO, ONTARIO M3J 1P3, CANADA; (NCC) DEPARTMENT OF ZOOLOGY, ERINDALE CAMPUS, UNIVERSITY

OF

TORONTO,

MISSISSAUGA,

ONTARIO L5L 1C6, CANADA. Send reprint re-

quests to DJMcQ. Submitted: 9 March 1995. Accepted: Ross.

6 Dec. 1995. Section editor: S. T.
