Morphological correlates of swimming activity in wild largemouth bass (Micropterus salmoides) in their natural environment

Individual variation in morphology has been linked to organismal performance in numerous taxa. Recently, the relationship between functional morphology and swimming performance in teleost fishes has been studied in laboratory experiments. In this study, we evaluate the relationship between morphology and swimming activity of wild largemouth bass (Micropterus salmoides) during the reproductive period, providing the first data derived on free-swimming fish not exposed to forced swim trials in the laboratory. Sixteen male largemouth bass were angled from their nests, telemetered, and subsequently monitored by a whole-lake acoustic hydrophone array with sub-meter accuracy. Additionally, eleven morphological measurements were taken from digital images of each fish. A principal components analysis of the morphological measurements described 79.8% of the variance. PC1 was characterized by measures of overall body stoutness, PC2 was characterized by measures of the length and depth of the caudal region, and PC3 characterized individuals with relatively large anterior portions of the body and relatively small caudal areas. Of these variables, only PC3 showed significant relationships to swimming activity throughout the parental care period. PC3 was negatively correlated with multiple measures of swimming activity across the parental care period. Furthermore, swimming performance of individual male bass was noted to be repeatable across the parental care period indicating that this phenomenon extends beyond the laboratory.     

Density-dependent effects of prey defences

In this study, we show that the protective advantage of a defence depends on prey density. For our investigations, we used the predator-prey model system Chaoborus-Daphnia pulex. The prey, D. pulex, forms neckteeth as an inducible defence against chaoborid predators. This morphological response effectively reduces predator attack efficiency, i.e. number of successful attacks divided by total number of attacks. We found that neckteeth-defended prey suffered a distinctly lower predation rate (prey uptake per unit time) at low prey densities. The advantage of this defence decreased with increasing prey density. We expect this pattern to be general when a defence reduces predator success rate, i.e. when a defence reduces encounter rate, probability of detection, probability of attack, or efficiency of attack. In addition, we experimentally simulated the effects of defences which increase predator digestion time by using different sizes of Daphnia with equal vulnerabilities. This type of defence had opposite density-dependent effects: here, the relative advantage of defended prey increased with prey density. We expect this pattern to be general for defences which increase predator handling time, i.e. defences which increase attacking time, eating time, or digestion time. Many defences will have effects on both predator success rate and handling time. For these defences, the predator’s functional response should be decreased over the whole range of prey densities. Received: 15 September 1999 / Accepted: 23 December 1999     

Effects of 17-beta estradiol and 4-nonylphenol on phase II electrophilic detoxification pathways in largemouth bass (Micropterus salmoides) liver

The effects of in vivo exposure to a natural and synthetic estrogen upon three hepatic phase II enzyme pathways involved in cellular protection against reactive intermediates were investigated in the largemouth bass (Micropterus salmoides). The pathways analyzed included glutathione S-transferases (GST), glutathione (GSH) biosynthesis and NAD(P)H-dependent quinone reductase (QR). Following exposure to 17-beta estradiol (E2, a model natural estrogen; 2 mg/kg, i.p.) or 4-nonylphenol (NP, a model synthetic estrogen; 5 mg/kg and 50 mg/kg, i.p.), serum vitellogenin concentrations in male fish were markedly increased. Exposure to E2 did not affect steady-state GST-A mRNA expression, although GST catalytic activity toward 1-chloro 2,4-dinitrobenzene (CDNB) was elevated at 48 h post-injection. In addition, the rates of bass liver GST-4-hydroxy-2-nonenal (GST-4HNE) conjugation were elevated by E2 exposure at all timepoints. In contrast, exposure to NP decreased steady-state GST-A mRNA levels, but did not alter GST catalytic activities. Hepatic GSH levels were not significantly affected by exposure to either compound, although a trend towards increased GSH biosynthesis was observed with both compounds. Although bass liver quinone reductase catalyzed 2,6-dichloroindophenol (DCP) reduction, unlike in rodents, these catalytic activities were not inhibited by dicoumarol. Exposure to 5 mg/kg NP significantly increased hepatic QR activities. Collectively, our data suggest that exposure to E2 or NP alters the ability of largemouth bass to biotransform environmental chemicals through glutathione S-transferase and quinone reductase catalytic pathways.     

Density-dependent foraging in the harvester ant Messor ebeninus: two experiments

Harvester ants are important seed predators in many xeric environments, and their foraging choices can influence the composition of plant communities. Seed abundance has been cited as an important factor in determining such foraging preferences. Three seed types (sesame, millet, and flax) were experimentally introduced in differing proportions near nests of the ant Messor ebeninus near the Dead Sea, in territory administered by the state of Israel. Two experiments were designed to investigate the effects of this density conditioning on two stages of the ants' subsequent foraging behavior: recruitment to seed patches and selection of seeds from within a patch. When seeds were presented in small, single-species patches, experimentally common seeds were discovered and exploited significantly faster than rare seeds, especially among less preferred seed types. When seeds were presented in large, mixed patches, however, no consistent effect of density was observed.     

Density-dependent effects of multiple predators sharing a common prey in an endophytic habitat

Multiple predator species feeding on a common prey can lead to higher or lower predation than would be expected by simply combining their individual effects. Such emergent multiple predator effects may be especially prevalent if predators share feeding habitat. Despite the prevalence of endophagous insects, no studies have examined how multiple predators sharing an endophytic habitat affect prey or predator reproduction. We investigated density-dependent predation of Thanasimus dubius (Coleoptera: Cleridae) and Platysoma cylindrica (Coleoptera: Histeridae) on a bark beetle prey, Ips pini (Coleoptera: Scolytidae), in a laboratory assay. I. pini utilize aggregation pheromones to group-colonize and reproduce within the stems of conifers. T. dubius and P. cylindrica exploit these aggregation pheromones to arrive simultaneously with the herbivore. Adult T. dubius prey exophytically, while P. cylindrica adults enter and prey within the bark beetle galleries. Larvae of both predators prey endophytically. We used a multiple regression analysis, which avoids confounding predator composition with density, to examine the effects of varying predator densities alone and in combination on herbivore establishment, herbivore reproduction, and predator reproduction. Predators reduced colonization success by both sexes, and decreased I. pini reproduction on a per male and per female basis. The combined effects of these predators did not enhance or reduce prey establishment or reproduction in unexpected manners, and these predators were entirely substitutable. The herbivores net replacement rate was never reduced significantly below one at prey and predator densities emulating field conditions. Similar numbers of each predator species emerged from the logs, but predator reproduction suffered from high intraspecific interference. The net replacement rate of P. cylindrica was not affected by conspecifics or T. dubius. In contrast, the net replacement rate of T. dubius decreased with the presence of conspecifics or P. cylindrica. Combinations of both predators led to an emergent effect, a slightly increased net replacement rate of T. dubius. This may have been due to predation by larval T. dubius on pupal P. cylindrica, as P. cylindrica develops more rapidly than T. dubius within this shared habitat.     

Context-dependent effects of bluegill in experimental mesocosm communities

Most knowledge of direct and indirect effects of zooplanktivorous fish has come from studies in which a treatment with a zooplanktivore is compared to a fishless control. However, effects of a zooplanktivore may be different in the presence of other fish species because the other fish have direct and indirect effects that may alter the effects of the zooplanktivore in question. To test this hypothesis, we conducted a tank mesocosm experiment of 2×2 factorial design in which the presence and absence of bluegill (Lepomis macrochirus) were cross-classified with the presence and absence of a fish assemblage composed of western mosquitofish (Gambusia affinis) and common carp (Cyprinus carpio). The presence of bluegill decreased Daphnia, Ceriodaphnia, cyclopoid copepodids, calanoid copepodids, copepod nauplii, amphipods, gastropods, and notonectids. Daphnia, Ceriodaphnia, cyclopoid copepodids, copepod nauplii, gastropods, notonectids, Najas, and Chara were decreased and herbivorous rotifers, turbidity, chlorophyll a, total nitrogen and total phosphorus were increased in the presence of the fish assemblage. Significant bluegill×fish assemblage interaction effects were detected for Daphnia, Ceriodaphnia, cyclopoid copepodids, copepod nauplii, gastropods, and notonectids. Analysis of simple effects for these response variables revealed that all significant bluegill effects in the absence of the fish assemblage were not significant in the presence of the fish assemblage. Our results indicate that the effects of bluegill may be context dependent, or dependent upon the presence of other trophically similar fish species. Received: 3 November 1998 / Accepted: 24 September 1999     

Identification of centrarchid hepcidins and evidence that 17β-estradiol disrupts constitutive expression of hepcidin-1 and inducible expression of hepcidin-2 in largemouth bass (Micropterus salmoides)

Hepcidin is a highly conserved antimicrobial peptide and iron-regulatory hormone. Here, we identify two hepcidin genes (hep-1 and hep-2) in largemouth bass (Micropterus salmoides) and smallmouth bass (Micropterus dolomieu). Hepcidin-1 contains a putative ATCUN metal-binding site in the amino-terminus that is missing in hepcidin-2, suggesting that hepcidin-1 may function as an iron-regulatory hormone. Both hepcidins are predominately expressed in the liver of largemouth bass, similar to other fish and mammals. Experimental exposure of pond-raised largemouth bass to 17β-estradiol and/or the bacteria Edwardsiella ictaluri led to distinct changes in expression of hep-1 and hep-2. Estradiol reduced the constitutive expression of hep-1 in the liver. Bacterial exposure induced expression of hep-2, suggesting that hepcidin-2 may have an antimicrobial function, and this induction was abolished by estradiol. To our knowledge, this is the first report of the regulation of hepcidin expression by estradiol in either fish or mammals.     

Effects of crayfish size,orientation, and movement on the reactive distance of largemouth bass foraging in clear and turbid water

Laboratory experiments were performed in clear and turbid water to determine the effects of prey size, orientation, and movement on the reactive distance of largemouth bass (Micropterus salmoides) when feeding on crayfish (Procambarus acutus). In clear water, the reactive distance increased linearly with an increase in prey size, and prey movement resulted in a significant increase in the reactive distance. Prey orientation (head-on versus perpendicular) did not change the reactive distances. In moderately turbid water, the reactive distance did not increase with increased prey size, and prey movement did not result in any changes in the reactive distance. The absence of any effects of prey orientation in clear water or prey movement in turbid water is inconsistent with results from studies using different species (primarily planktivorous fish). I propose that largemouth bass change their foraging tactics as prey visibility changes. When prey are highly visible (low turbidity), predators attack (react) only after prey recognition, which is based on multiple cues such as prey size (length, width) and movement. When prey are less visible (high turbidity), predators attack immediately upon initial prey sighting, which does not depend on prey size or movement.     

Sex and individual differences in foraging behavior of Japanese cormorants in years of different prey availability

Japanese cormorants, Phalacrocorax capillatus, are sexually dimorphic seabirds with males that are heavier and that dive deeper than females. Sex differences in prey composition and foraging behavior of those rearing chicks at Teuri Island, Hokkaido, were examined by collecting food-loads from parents in 1992–1998 and by radio-tracking ten birds each in 1997 and 1998 when prey availability was different. Males fed more on benthic and epibenthic fishes (82% mass) than did females (34%) while females fed more on epipelagic and coastal fishes (53%) than did males (18%). Males made longer dives (53 s) at feeding sites closer to the island (7 km) than females (39 s, 13 km) in 1997. In 1998 when the availability of epipelagic fish seemed to be higher, there were no sex differences in dive duration and distance to the feeding sites (35 s and 9 km for males, 36 s and 10 km for females). This sex difference in foraging behavior with a poor availability of epipelagic fish suggests that high diving ability possibly enables males to feed on demersal fish. Birds specializing in coastal shallow waters around the island made long dives; hence they were probably foraging in bottom layers. Those foraging in deeper shelf waters made short dives and they were thought to forage in surface layers. Electronic Publication     

The diet of largemouth bass, Micropterus salmoides, in Lake Naivasha, Kenya

Lake Naivasha is a freshwater lake situated in the eastern rift valley of Kenya. Only five species of fish are present, all of which have been introduced. Of these, Oreochromis leucostictus, Tilapia zillii and Micropterus salmoides (largemouth black bass) support an important gillnet fishery with bass also being taken for sport. Until bass reached 260 mm f.l. they depended upon invertebrate food organisms. Thereafter crayfish, fish and frogs became increasingly important the larger the size of the bass. The most important invertebrate prey species was the water boatman, Micronecta scutellaris , followed by chironomid and culicid pupae. Zooplank-ton was consumed but only in large quantity by fish smaller than 80 mm. For bass over 260 mm the crayfish, Procambarus darkii , was the principal food. The largemouth bass in Lake Naivasha are generalized macro-predators, feeding principally on free-living animals of a kind most likely to be found in the littoral zones.     

Overcompensatory recruitment and generation delay in discrete age-structured population models

 The effect of overcompensatory recruitment and the combined effect of overcompensatory recruitment and generation delay in discrete nonlinear age-structured population models is studied. Considering overcompensatory recruitment alone, we present formal proofs of the supercritical nature of bifurcations (both flip and Hopf) as well as an extensive analysis of dynamics in unstable parameter regions. One important finding here is that in case of small and moderate year to year survival probabilities there are large regions in parameter space where the qualitative behaviour found in a general n+1 dimensional model is retained already in a one-dimensional model. Another result is that the dynamics at or near the boundary of parameter space may be very complicated. Generally, generation delay is found to act as a destabilizing effect but its effect on dynamics is by no means unique. The most profound effect occurs in the n-generation delay cases. In these cases there is no stable equilibrium X ^* at all, but whenever X ^* small, a stable cycle of period n+1 where the periodic points in the cycle are on a very special form. In other cases generation delay does not alter the dynamics in any substantial way. Received 25 April 1995; received in revised form 21 November 1995     

Stochastic density dependence in population size of a benthic clonal invertebrate: the regulating role of fission

The influence of environmental variation on the demography of clonal organisms has been poorly studied. I utilise a matrix model of the population dynamics of the intertidal zoanthid Palythoa caesia to examine how density dependence and temporal variation in demographic rates interact in regulating population size. The model produces realistic simulations of population size, with erratic fluctuations between soft lower and upper boundaries of approximately 55 and 90% cover. Cover never exceeds the maximum possible of 100%, and the population never goes to extinction. A sensitivity analysis indicates that the model’s behaviour is driven by density dependence in the fission of large colonies to produce intermediate sized colonies. Importantly, there is no density-dependent mortality in the model, and density dependence in recruitment, while present, is unimportant. Thus it appears that the main demographic processes which are considered to regulate population size in aclonal organisms may not be important for clonal species. Received: 18 August 1999 / Accepted: 29 October 1999     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000     

Nutritional condition and physiology of paternal care in two congeneric species of black bass (<Emphasis Type="Italic">Micropterus</Emphasis> spp.) relative to stage of offspring development

Parental care requires a complex integration of physiology and behaviour, yet little is known about the physiological and energetic consequences or correlates of these behaviours. Using two species of male black bass (smallmouth bass, Micropterus dolomieu; largemouth bass, M. salmoides) as a model, the focus of this study was to determine the biochemical and hematological indicators of change in nutritional status and potential for chronic stress. This was accomplished by randomly sampling individuals at four stages across parental care. Additionally, a subset of individuals was repeatedly sampled at three brood development stages to track changes in biochemical factors within the individual. Though there were changes in physiological factors across parental care in randomly sampled fish of both species (declines in plasma glucose in largemouth bass; decreases in hematocrit and plasma chloride in smallmouth bass), repeated sampling of individuals was determined to be a more appropriate sampling technique due to natural variability in biochemical factors among individual fish. Repeated sampling of smallmouth bass did not adversely influence physiological metrics or brood abandonment. However, there were higher incidences of nest abandonment in repeatedly sampled largemouth bass. Amongst the repeatedly sampled smallmouth bass, nutritional indicators such as plasma triglyceride levels decreased indicating individual fasting across the majority of parental care. Increases in plasma calcium and magnesium towards the end of care indicated that feeding most likely resumed when the brood was close to independence after ~3 weeks of care. Lastly, several indicators of chronic stress, such as plasma glucose and chloride levels, increased throughout the parental care period. These sublethal stressors are indicative of decreasing body condition associated with prolonged activity and fasting which may have marked impacts on the ability of an individual to continue parental care for the current brood and impact subsequent individual fitness. Further research into the mechanistic relationships between behaviour, physiology, and energetics during the parental care period will provide a better understanding of the decisions by individuals facing multiple trade-offs that ultimately lead to differences in individual fitness.     

Interspecific competition among macroparasites in a density-dependent host population

 We analyze the dynamics of a community of macroparasite species that share the same host. Our work extends an earlier framework for a host species that would grow exponentially in the absence of parasitism, to one where an uninfected host population is regulated by factors other than parasites. The model consists of one differential equation for each parasite species and a single density-dependent nonlinear equation for the host. We assume that each parasite species has a negative binomial distribution within the host and there is zero covariance between the species (exploitation competition). New threshold conditions on model parameters for the coexistence and competitive exclusion of parasite species are derived via invadibility and stability analysis of corresponding equilibria. The main finding is that the community of parasite species coexisting at the stable equilibrium is obtained by ranking the species according t! o th e minimum host density H ^* above which a parasite species can grow when rare: the lower H ^* , the higher the competitive ability. We also show that ranking according to the basic reproduction number Q ₀ does not in general coincide with ranking according to H ^* . The second result is that the type of interaction between host and parasites is crucial in determining the competitive success of a parasite species, because frequency-dependent transmission of free-living stages enhances the invading ability of a parasite species while density-dependent transmission makes a parasite very sensitive to other competing species. Finally, we show that density dependence in the host population entails a simplification of the portrait of possible outcomes with respect to previous studies, because all the cases resulting in the exponential growth of host and parasite populations are eliminated.. Received: 24 June 1996 / Revised version: 28 April 1998     

Influence of prey availability and conspecifics on patch quality for a cannibalistic forager: laboratory experiments with the wolf spider Schizocosa

 Because cannibals are potentially both predator and prey, the presence of conspecifics and alternative prey may act together to influence the rate at which cannibals prey upon each other or emigrate from a habitat patch. Wolf spiders (Lycosidae) are cannibalistic-generalist predators that hunt for prey with a sit-and-wait strategy characterized by changes in foraging site. Little information is available on how both prey abundance and the presence of conspecifics influence patch quality for these cursorial, non-web-building spiders. To address this question, laboratory experiments were conducted with spiderlings and older juveniles of the lycosid genus Schizocosa. The presence of insect prey consistently reduced rates of spider emigration when spiders were housed either alone or in groups. Solitary juvenile Schizocosa that had been recently collected from the field exhibited a median giving-up time (GUT) of 10 h in the absence of prey (Collembola); providing Collembola increased the median GUT to 64 h. For solitary spiders, the absence of prey increased by about fourfold the rate of emigration during the first 24 h. In contrast, for spiders in patches with a high density of conspecifics, the absence of prey increased the 24-h emigration rate by only 1.6-fold. For successful cannibals in the no-prey patches, the presence of conspecifics improved patch quality by providing a source of food. Mortality by cannibalism was affected by both prey availability and openness of the patch to net emigration. In patches with no net emigration, the presence of prey reduced rates of cannibalism from 79% to 57%. Spiders in patches open to emigration but not immigration experienced a rate of cannibalism (16%) that was independent of prey availability. The results of these experiments indicate that for a cannibalistic forager such as the wolf spider Schizocosa, (1) the presence of conspecifics can improve average patch quality when prey are absent, and (2) cannibalism has the potential to be a significant mortality factor under natural field conditions because cannibalism persisted in prey patches that were open to emigration. Received: 12 April 1996 / Accepted: 14 August 1996     

Linking individual foraging behavior and population spatial distribution in patchy environments: a field example with Mediterranean blue tits

 In spite of recent theoretical interest, few field studies have addressed the links between individual behavioral decisions and population distribution. This work analyzes the foraging behavior of individuals and the spatial distribution of a population of blue tits (Parus caeruleus) just before the main breeding season, when blue tit foraging was not affected by central-place or flocking behaviors. The study was carried out in open holm oak Quercus ilex woodlands (dehesas) that are patchy for canopy-foraging birds because of the scattered arrangement of trees. Residence time on each tree was not correlated either with previous flight time or with prey abundance in trees. Flight distances between trees were larger than average distances estimated in random samples of holm oaks taken close to foraging birds. Trees were not selected by birds on the basis of their expected energy costs and rewards. Bird abundance was not related to food availability in trees or to tree size across dehesas. However, bird abundance was strongly correlated with tree density and with the availability of tree holes for nesting, to the extent that the proportion of tits matched the proportions of both tree abundance and hole abundance across study plots. Overall, neither the behavior of individuals nor the distribution of the population of blue tits corresponded with food resources, which appeared superabundant; instead, tits appeared to behave and be distributed according to the distribution of structural resources such as trees and tree holes for nesting. Received: 18 August 1996 / Accepted: 14 March 1997     

Predation of Japanese dace, Tribolodon hakonensis, by largemouth bass, Micropterus salmoides, in experimental aquaria

 To test the size range of prey fish that largemouth bass, Micropterus salmoides, can successfully consume, live Japanese dace, Tribolodon hakonensis, were given as prey fish to individual largemouth bass in aquaria. The ratio of maximum standard length (SL) of the Japanese dace consumed by largemouth bass was 46–69% of bass SL. The maximum length of Japanese dace consumed did not differ significantly between largemouth bass and smallmouth bass (M. dolomieu) previously studied, although largemouth bass have relatively larger mouth sizes than smallmouth bass. Largemouth bass occasionally injured and killed Japanese dace larger than the limit that could be consumed.     

Diet overlap among ruminants in Fennoscandia

Information on overlap in resource use is central to understanding of interspecific exploitation competition and resource partitioning. Despite this, measures of diet overlap among northern ruminants in Fennoscandia is limited to one earlier study (reindeer and sheep). Diet overlap between sympatric moose and roe deer calculated with Schoener’s index was 20.7% and 33.6% during summer (data from one area) and winter (data from two areas), respectively, whereas average diet overlap between moose and red deer was 32.0% during winter (data from four areas). Diet overlap between a coastal island population of red deer and sheep was 59.3% during summer and 63.9% during winter. Summer diet overlap between a sheep and a goat population and a sheep and a reindeer population calculated with data on main types of forage plants was 77.0% and 55.1%, respectively. However, overlap calculated with main plant groups was sometimes considerably higher than when calculated for individual forage species. Neither difference in feeding type nor body mass successfully predicted diet overlap between species pairs (n=9), although there tended to be negative correlation (r _p =–0.586, P=0.098) between diet overlap of main plant groups (calculated across studies) and difference in feeding type. Received: 19 October 1999 / Accepted: 31 January 2000     

Density-dependent variation in reproductive success in a terrestrial orchid

 Within population variation in plant density can affect reproductive success and breeding systems. We examined such effects in Listera cordata R. Br., a north temperate terrestrial orchid, in a subalpine population in Colorado, USA. Hand pollinations showed that L. cordata was self-compatible as all pollinations produced fruits. Cross-pollinations differed significantly from self-pollinations, and had a higher number of seeds per capsule and higher % of fertilized ovules. Together they could compensate for the transmission advantage of self-pollinations. Average number of flowers per inflorescence was 4.2–4.4 and average fruit set was 20.7–23.7%. By establishing 36 plots with varying plant densities (1, 4, 8 and 16) and emasculating all flowers in target plants of a set of 20 plots, we estimated density effects on fruit and seed production. Plant density had no effect on the amount of reproduction except in reduced seed production of highly dispersed plants. Our results suggest that inbreeding depression may be an important selective factor in L. cordata. Further studies are needed to define its role in the evolution of mating systems in this species. Received February 21, 2000 Accepted December 26, 2000