Population genetic structure of walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> (Gadidae,Pisces) from the Bering Sea and Sea of Okhotsk

Walleye pollock Theragra chalcogramma Pallas occupies a central place in ecosystems of the North Pacific and is an important target species of fisheries. The species is characterized by daily vertical, spawning, feeding, and wintering migrations and spawning occurring under the sea ice. Since population structure estimation by the tagging with recapture is inefficient in walleye pollock, the pollock resources are difficult to estimate by conventional methods, requiring population genetic studies with molecular markers. The population genetic structure of five spawning aggregations from the Bering Sea was for the first time studied with ten microsatellite loci: Tch5, Tch10, Tch11, Tch12, Tch14, Tch16, Tch17, Tch19, Tch20, and Tch22. A spatially distant sample from the Sea of Okhotsk was used as a reference group. Polymorphism for the markers reached 100%, and heterozygosity of individual loci ranged from 41 to 95% in different populations. It was shown the aggregations of interest are in goodness-to-fit the Hardy-Weinberg equilibrium (HWE) at hole, while the Sea of Okhotsk sample demonstrated a sex bias: the heterozygosity at Tch16 in males was significantly lower than in females. The highest discriminative power was observed for Tch10, Tch20, and Tch22. F _ST genetic distances between populations were typical for marine fishes. A mixed composition was supposed for the sample from the region of the underwater Shirshov Ridge, which serves as a natural partial geographic barrier between the Olyutor-Karagin and Koryak walleye pollock stocks. With the Shirshov sample excluded, F _ST scatter plots and the spatial autocorrelation approach supported isolation by distance for the aggregations. An influence of abiotic factors on the population structure was assumed for walleye pollock of the Bering Sea.     

Role of male-male competition and female choice in the development of breeding coloration in pupfish (Cyprinodon pecosensis)

The function of male breeding coloration and the conditionsunder which it is expressed were investigated using two experimentalprotocols that differed in the degree of control of social factors.In tank experiments, focal males were separated by opaque orclear partitions or allowed to interact freely without partitions.In lake experiments, males were introduced into opaque or clearenclosures in the lake from which they were collected. Thisdesign tested the relative importance of breeding substrate,social interactions with other males, and presence of femalesand juveniles on the expression of breeding coloration and agonisticand mating behavior in pupfish. Breeding coloration was notexpressed in the absence of conspecifics, even in the presenceof suitable breeding substrate, or in the presence of juveniles.Breeding coloration was poorly developed when only males werepresent, was more intense in the presence of females, and darkenedsignificantly after a spawning bout Males in clear enclosuresin their natural lake habitat were in visual contact with conspecificsof both sexes and developed more intense blue breeding colorationthan males in the tank experiments that were in visual contactonly with other males. Uncon-fined males with territories inthe lake developed the most intense breeding coloration. Breedingcoloration of male pupfish is a facultative trait that is welldeveloped when breeding females are present and plays a majorrole in the attraction and courtship of mates. However, maximumdevelopment of breeding coloration by territorial males suggeststhat intrasexual and intersexual selection act concordantlyon this trait.[Behav Ecol 7: 431-437 (1996)]     

Sex-biased egg cannibalism in spawning walleye pollock: the role of reproductive behavior

We studied inadvertent egg cannibalism in spawning stocks of walleye pollock, Theragra chalcogramma, in the Gulf of Alaska and the Bering Sea between 1986 and 1996. Male pollock had on average 3 times more eggs in their stomachs than females. For both sexes adult fish of average body length had more eggs in their stomach than did smaller or larger fish. When maturity of fish was taken into account, actively spawning males had the highest numbers of eggs in their stomachs. We found weak evidence for a diel variation; the number of eggs per stomach was high in both sexes during the day and lower during the night. We suggest two possible explanations for this phenomenon. Sex-biased egg cannibalism may reflect the differential time spent in layers of high egg densities. Hydroacoustic and trawl catch data from both areas suggest that males aggregate deeper than females. Spawning takes place in the deeper layers of the fish aggregation, so males spend more time in high egg densities. Alternatively, males may be more active than females and increased gill ventilation and/or drinking rates may be responsible for the differences in egg cannibalism.     

Influence of male aggregation size on female visitation in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae)

Abstract  The importance of male aggregation size for female visitation and initiation of male pheromone-calling was investigated in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) using artificial male aggregations in large laboratory cages. Female B. tryoni visited the largest aggregation more frequently than single males in association with a higher proportion of calling males, but there was no correlation between aggregation size and female visitation. Female B. tryoni had a limited capacity to perceive a difference between the number of calling males. Calling propensity of male B. tryoni was increased by the presence of conspecific males. Increased calling propensity in larger groups of male B. tryoni may be due to social facilitation of male calling behaviour. Female visitation at aggregations was only weakly associated with male calling, suggesting that aggregation size and the number of pheromone-calling males are not the only factors important in locating mates in B. tryoni , and it is possible that low-density populations could persist so long as females can encounter single males.     

Evaluation of a conservation strategy: a spawning aggregation closure for red hind, Epinephelus guttatus, in the U.S. Virgin Islands

Many tropical reef fishes spawn in large aggregations, which are readily targeted by fishers. By the 1980s, at least two grouper spawning aggregations were eliminated by intensive fishing off the island of St. Thomas, U.S. Virgin Islands, and another aggregating species, red hind, was intensively targeted. By 1988, the average length of red hind had greatly decreased to 295 mm, and the sex ratio was extremely skewed to 15 females per male, suggesting a heavily fished stock. Since this species is a protogynous hermaphrodite, the loss of large individuals (primarily males) could potentially result in sperm limitation in spawning aggregations. In 1990, a spawning aggregation closure was implemented. By 1997, average size of red hind had increased to 395 mm and sex ratio had shifted to 4 females per male. Fish were observed aggregating only in structurally complex habitat along the insular shelf edge. This habitat type is apparently not common along most of the shelf edge off St. Thomas and may provide shelter while reducing risk of predation during aggregation periods. These data suggest that protection of spawning aggregations is a sound management strategy with considerable potential for aiding the sustainable use of reef fish resources.     

Aggregation and spawning of the humphead wrasse Cheilinus undulatus (Pisces: Labridae): general aspects of spawning behaviour

The humphead wrasse Cheilinus undulatus formed resident spawning aggregations daily after high tide at specific locations along the seaward edge of the Palau barrier reef. The location and extent of one aggregation site remained consistent for 6 years with no physical features distinguishing it from adjacent areas. Spawning was documented most months and probably occurred year round with possible seasonal and lunar variation. Spawning males arrived first at the site, followed by females and potentially small primary males. The aggregation female to male sex ratio was estimated to be between 6:1 and 10:1. A maximum of 15 males and 100–150 females were observed at the site. A male courtship posture with the anal fin pointed, the caudal fin folded down and the dorsal fin folded against the body was maintained while swimming a few metres off the bottom in view of females. When ready to spawn females rose up as the posturing male passed and the pair released gametes in a relatively sedate fashion near the surface along the shelf break. No attempted predation on spawning adults was seen. Egg predation after spawning was uncommon. On days with early to mid‐day high tides the spawning period started 2·0–2·5 h after high tide when the speed of lagoon–ocean tidal currents peaked and lasted c. 1 h. On days with later afternoon high tides, spawning occurred sooner after high tide and before current speeds peaked. Other fishes with planktonic eggs spawned at the site as pairs or small groups in a rough succession after high tide with C. undulatus, the last species to spawn.     

The biochemical composition and calorie density of the walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> in the Sea of Okhotsk

In tissues of the walleye pollock Theragra chalcogramma the dry matter content averages 18.5%. The lipid content of the raw material is 0.7%, the protein content is 15.3%, carbohydrates are 0.6%, and ash is 1.3%. The average calorie density is 940 cal/g wet weight and 5080 cal/g dry weight. The dry matter content of gonads varies within 14.9–28.0% in females and 14.5–17.0% in males. The lipid content of the raw material is 0.9–3.0% in females and 1.3–1.8% in males; the protein content is 10.2–21.5% and 10.7–13.4%, respectively. The calorie density of female gonads is 702–1537 cal/g wet weight and 4426–5482 cal/g dry weight; for the male gonads it is 760–960 cal/g wet weight and 4952–5641 cal/g dry weight. The dry matter content of the liver varies within 42.2–62.2% for females and 34.4–62.4 for males. The lipid content of the raw material is 25.6–44.5% for females and 16.6–41.3% for males; the protein content is 6.3–9.8% and 8.1–12.3%, respectively. The calorie density of the liver in females varies within 2918–4601 cal/g wet weight and 6370–7395 cal/g dry weight; in males it is 2291–4357 cal/g wet weight and 6392–7492 cal/g dry weight. The minimum calorie density of the liver is observed in juvenile pollock: 963 cal/g wet weight and 2045 cal/g dry weight. The dry-matter content of feces in different size groups varies within 15.0–18.4%. Values of the average lipid content of raw material range from 1.1 to 1.6%; the protein content is from 1.8 to 3.8% and carbohydrates are from 0.9 to 1.4%. The calorie density of feces from variously-sized walleye pollock varies within a narrow range, from 308 to 362 cal/g wet weight. The energy equivalent ranges, depending on body size, from 259 to 2377 cal. The share of energy concentrated in the somatic (muscle) tissue of variously-sized walleye pollock during ontogenesis constitutes 56.5–93.9%; in female gonads it is 0.9–26.6%; in male gonads it is 0.4–7.3%, in the female liver it is 7.9–27.2%, and in the male liver it is 5.7–26.9%. The amount of energy (cal), concentrated in the female liver and gonads is on average 1.5 and 3 times as high as that in the male liver and gonads, respectively. The maximum total energy loss (15–30%) in mature walleye pollock of various-sizes occurs in the spawning period, during the transition from the maturity stage 5 to stage 6. The total amount of energy accumulated during the lifecycle from small juveniles (<17 cm) to very large individuals (>60 cm) averages 1964 kcal for females and 1465 kcal for males. The difference in the amount of energy is explained by the fact that oogenesis requires more energy than spermatogenesis.     

Spawning behavior of the tiger grouper (<Emphasis Type="Italic">Mycteroperca tigris</Emphasis>) in a Caribbean atoll

Many grouper (Epinephelidae) species aggregate to spawn at specific times of year, at consistent locations, and at specific times of day and phases of the moon. The tiger grouper (Mycteroperca tigris) is widely distributed in the Caribbean Sea and aggregates to spawn just after the full moon. In January and February 2003–2005, we conducted visual surveys of a tiger grouper spawning aggregation at Glover’s Reef, Belize. The primary objectives of this study were to describe grouper spawning activities with respect to the timing of the lunar cycle, identify and describe male and female spawning behaviors, and estimate the number of grouper spawning events each day. The peak number of both male and female tiger groupers counted each year occurred from 6 to 10 d after the full moon. The peak count of males was 1.6–2.2 times greater than the peak count of females. Six categories of male–male and six categories of male–female interactions were identified. All male tiger groupers displayed an ability to rapidly change color; this was often triggered by a close encounter with other male or female tiger groupers. The daily spawning activity was remarkably predictable. The first spawning occurred on average 8.1 min before sunset, and the last occurred 5.1 min after sunset. On average, less than half of the females at the spawning site spawned each day, but some females spawned more than once in a given day. Females appeared to choose their mates.     

Effect of Population Density,Size of Habitat and Oviposition Substrate on the Breeding System of Pupfish (Cyprinodon pecosensis)

Size of habitat, availability of oviposition substrate, and population density were manipulated to determine their effects on the structure and dynamics of the pupfish breeding system. The fish established a territorial breeding system in large tanks, irrespective of population density or oviposition substrate. A dominance hierarchy, in which one male controlled most of the oviposition substrate and spawned with most females, was established in small tanks at low densities. Both population density and oviposition substrate affected the number of males defending territories. Male spawning success was inversely related to availability of oviposition substrate, but was not affected either by density or tank size. Location of a territory and its size also influenced spawning success. Males with bigger territories had higher spawning success. A “carryover” effect of neighbors was observed; males with reproductively active neighbors spawned more often than those with less successful neighbors. Interference with spawning was affected both by tank size and density. Agonistic behavior of breeding males was not affected by any manipulation, but it was positively associated with male spawning success at high but not at low densities. In all treatments the mortality of breeding males was approximately 8 times as high as that of females, suggesting that, under these experimental conditions, the cost of reproducing is substantially higher for males. Comparisons between breeding pupfish in these experiments and in natural environments provide a basis for understanding the selective pressures that have favored the maintenance of these extremely plastic breeding systems.     

Sexual reproduction in the tropical corallimorpharian Rhodactis rhodostoma

Abstract. Polyps of the tropical corallimorpharian Rhodactis rhodostoma segregate sexes between center and edge positions within aggregations produced by clonal replication. On a reef flat at Eilat, northern Red Sea, infertile polyps and males occur mainly along the edges of clonal aggregations, while females mostly occupy central positions within each aggregation. In addition, on the inner to middle reef flat where polyps of this species are abundant, aggregations consist mostly of females. On the outer reef flat, where polyps are rare, a sampled aggregation consisted mostly of males and infertile polyps. Male polyps are significantly smaller than females, and the smallest polyps are infertile. Fecundity increases significantly with polyp size in females, but testis size and number do not vary with body size in males. Oocytes are present in polyps during most of the year and gradually increase in size until annual spawning in June-July during the period of maximum day length. Testes do not vary significantly in size during the year and remain a small proportion of body mass (>8%). In contrast, females invest up to 30% of their body mass into gonads during the months immediately before spawning. The annual spawning of gametes coincides with a temporary drop in the frequency of clonal replication by polyps. We estimate that each female polyp of R. rhodostoma may release up to 3000 large eggs (500 μm in maximum diameter) each summer. The high investment of this corallimorpharian in sexual production of planktonic propagules may allow rapid dispersal to reef habitats distant from parent populations.     

Predation risk and the opportunity for female mate choice in a coral reef fish

The influence of predation risk on the opportunity for female mate choice was investigated in the tailspot wrasse Halichoeres melanurus at two sites on a coral reef in Okinawa, Japan. Females mated repeatedly with the nearest males, but they also changed mates frequently at both sites. Mate changes were seen not only in the context of spatiosocial changes (mate disappearance or shifts in male territories) but were also probably the result of actual mate choice by females. Females at one site (site A) changed mates more often and conducted longer spawning trips from their home ranges to male territories than at the other site (site B). Fish at site A were faced with a higher frequency of predators than that of site B. However, fish of site A suffered fewer attacks from predators because they had more shelter, suggesting lower predation risk in that site. These results suggest that females under higher predation risk had less opportunity to choose preferable mates and that they had to mate with the same, nearest males in most cases.     

Behavioral Tactics of Male Sockeye Salmon (Oncorhynchus nerka) under Varying Operational Sex Ratios

Previous studies have demonstrated several reproductive-behavior patterns in male salmon, including competitive and sneaking tactics, the formation of hierarchies, and non-hierarchical aggregations around ripe females. Through behavioral observations at varying spatial and temporal scales, we examined the hypothesis that operational sex ratio (OSR) determines male sockeye salmon (Oncorhynchus nerka) distribution and breeding tactics. Patterns of male distribution and behavior varied over both coarse and fine scales, associated with apparent shifts in reproductive opportunities, the physical characteristics of the breeding sites, and the deterioration of the fish as they approached death. Females spawned completely within a few days of arriving on the spawning grounds, whereas males courted the available ripe females from the date of their arrival on the spawning ground until their death. This difference in reproductive lifespans tended to elevate late-season OSRs but was partially counterbalanced by male departures and the arrival of other ripe females. The proportion of males able to dominate access to ripe females decreased and the number of large courting groups increased over the course of the season, apparently related to both increasing OSR and the deteriorating physical condition of males. However, great variation in OSR was observed within the spawning sites on a given day. OSRs were generally higher in shallow than in deep water, perhaps because larger females or more desirable breeding sites were concentrated in shallow water. The aggregations of males courting females were not stable (i.e. many arrivals and departures took place) and male aggression varied with group size. Aggression was most frequent at low OSRs and in groups of intermediate size (2–4 males per female), and much less frequent in larger groups, consistent with the needs of maximizing reproductive opportunities while minimizing unproductive energy expenditure. These results indicate that, while OSR strongly influences male distribution and behavior, other factors such as physical condition, limited movement and habitat choice may also affect male reproductive opportunities.     

Spawning of walleye pollock (<Emphasis Type="Italic">Theragra chalcogramma</Emphasis>) off the Southwestern coast of Kamchatka

The results of ichthyoplankton surveys conducted in 2007–2008 showed that waters off Southwest Kamchatka and North Kuril Islands were areas of mass spawning of walleye pollock. The peak spawning occurred during the last 10 days of April and in early May, which was much later than the peak at the main spawning site off West Kamchatka. The spawning activity of walleye pollock near the southwestern shores of Kamchatka is a regular event, as the analysis of archive materials shows. This gave us grounds to suspect the existence of a southern site that coincided well with the mass spawning in the Shelikhov Gulf in its timing and scale, but was missed during standard ichthyoplankton surveys conducted in early April. After analyzing the growth rates of spawners, the assumption was made that the southern spawning was performed by the East Kamchatkan walleye pollock population, whose mass spawning usually occurs in late April-early May. According to the data of 2008, the estimated biomass of walleye pollock spawning in the area of the Ozernovskaya basin in late April was nearly 600000 tons. Regular monitoring of the southern spawning is proposed by means of additional ichthyoplankton surveys south of 53° N, including the Okhotsk Sea waters of the North Kuril Islands, in late April—early May.     

Reversed Sex-Change in the Protogynous Reef Fish Labroides dimidiatus

Protogynous hermaphroditism, or female-to-male sex change, is known for many reef fishes including wrasses (family Labridae) in which large males monopolize mating. When the dominant male disappears from a polygynous group, the largest female may change sex within a few weeks. Such social control of sex change was first documented in harems of the cleaner wrasse Labroides dimidiatus almost 30 yr ago. To examine whether change of social status would induce males of L. dimidiatus to perform reversed sex-change, we conducted experiments: (i) releasing single males near lone males whose mates have been removed in the field; and (ii) keeping two males in a tank. Smaller males changed back to females when they became subordinate: it took 53–77 d (n=3) for them to complete gonadal sex change and release eggs in the aquarium. The male–male pairs performed spawning behavior, with the smaller male in the female role already 5–58 d before completion of gonadal sex change. This is the first report of reversed sex-change among protogynous wrasses. Moreover, we conducted another experiment, keeping a pair of a male and a larger female in a tank (n=1). We found sex change by both mates, which has not been reported from any fishes. Thus, the sex of L. dimidiatus is strictly determined by social status whenever it changes after mate loss.     

The influence of predator presence on utilization of artificial seagrass habitats by juvenile walleye pollock,Theragra chalcogramma

Synopsis Behavioral preference for a structured habitat (artificial seagrass) by juvenile walleye pollock,Theragra chalcogramma, was tested in controlled laboratory experiments. We monitored position of fish in 2000 1 tanks with and without artificial seagrass present in one half of the tank. In addition, we exposed walleye pollock to a predator model, assessing their response when a grass plot was available or unavailable as a potential refuge. In the absence of predators, the fish avoided the artificial seagrass, displaying a preference for the open water side of the experimental tanks. In the presence of a predator model, however, juvenile walleye pollock readily entered the artificial seagrass plots. In addition, they often remained in the grass canopy in proximity to the predator instead of moving out of the grass to avoid the predator (when no grass was present they consistently moved to the opposite side of the tank from the predator). The behavioral choices exhibited in this study suggest that juvenile walleye pollock modify habitat selection in response to perceived predation risk, and recognize the structure provided by artificial seagrass as a potential refuge.     

Reproductive behavior and mate fidelity in the monogamous goby,Valenciennea longipinnis

Reproductive behavior and mate fidelity of the gobiid fish,Valenciennea longipinnis, were studied on the coral reef at Sesoko Island, Okinawa, Japan. These fish usually live in pairs, not only foraging together for benthic animals in sandy areas, but also constructing several burrows within their home range. Before spawning, both fish, although mainly the male, constructed a mound, piling up dead-coral fragments, pebbles, shells, sand and algae onto one of the burrows. After spawning an egg mass on the ceiling of the burrow, the female stayed outside and continued the construction and maintenance of the mound for 3–5 days until hatching, while the male tended the eggs inside. Mate guarding of females seemed to prevent males from monopolizing several females. Although some pairs showed mate fidelity through several spawnings, more than half of the pairs broke up after only one spawning. The pair bond was broken by mate desertion and the disappearance of each sex. Both sexes preferred larger spawning partners; larger females spawned more eggs and larger males provided better egg care. Mate desertion occurred when larger potential mates, relative to the current partner, became available. The frequency of solitary individuals was higher in males than in females, resulting in females deserting their mates more often than males. Two factors seem to have facilitated mate desertion: (1) occurrence of size mis-matched pairing and (2) overlapping home ranges.     

Seasonal changes in energy and the energy cost of spawning in Gulf of Alaska Pacific cod

In the Gulf of Alaska, adult Pacific cod exhibited an annual cycle of condition, gonad index and liver index in which maximum values occurred in ripe fish in March and minima in July. About 30–31 % of prespawning stored energy was expended during the spawning effort. The energy associated with spawning derived from liver (24% and 18%), somatic tissue (22% and 33%) and gonad (53% and 48%) for females and males, respectively. Liver index and gonad index at the time of sampling were directly related in females, but in males gonad index was best related to liver index 1–3 months earlier.
The Pacific cod is very similar to the Atlantic cod in terms of energy cycling, maximum gonad sizes, energy expended during spawning and gonadal contribution to energy expenditure. However, in Pacific cod, somatic tissue contributes markedly to energy expended during reproduction. The Pacific cod cod differs from the walleye pollock with respect to gonad index (13% and 20%ν. 20% and 8% for females and males, respectively), spawning weight loss (25%ν. 38%), liver energy loss during spawning (71%ν. 55%) and energy cost of spawning.     

Interannual differences in growth and hatch date distributions of age-0 year walleye pollock Theragra chalcogramma (Pallas) sampled from the Shumagin Islands region of the Gulf of Alaska, 1985 – 2001

Daily increments of age-0 year walleye pollock Theragra chalcogramma otoliths from 1985 to 2001 were examined for interannual variability in growth and hatch dates. Fish were collected in summer and autumn surveys near the Shumagin Islands in the western Gulf of Alaska (GOA). Hatch date distributions of these fish were compared with hatch dates of larvae from spawning aggregations located in Shelikof Strait and the Shumagin Islands based on their spawning times. The hatch date distributions of age-0 year fish captured from the Shumagin Islands area were similar to those observed for larvae from the Shelikof Strait spawning group. Age-0 year fish whose hatch dates corresponded to the Shumagin Islands spawning, which occurred earlier in January and February, were not found. Sea surface temperature was associated with variability in hatch date distribution and growth.     

Female clustering in cockroach aggregations—A case of social niche construction?

Individuals in groups can suffer costs through interactions with adversarial or unknown conspecifics. Social niche construction allows individuals to buffer such potential costs by only engaging in preferred associations. This may be particularly beneficial in insect aggregations, which are often large and highly fluid. However, little is known regarding the structuring of such aggregations. Here we use social network analyses to test for fine‐scale social structure in resting aggregations of the sub‐social cockroach Diploptera punctata and to explore the social pressures that contribute towards such structure. We showed that females were significantly more gregarious than males and formed the core of the proximity network, thus demonstrating a higher level of social integration. This fine‐scale structure is likely to result from females displacing males; females initiated most displacements whilst males received the majority. We explain this behaviour in terms of social niche construction by showing that females received significantly fewer approaches and investigations at more female‐biased local sex ratios. We therefore suggest that female social clustering occurs in this, and presumably other, species to reduce potential costs associated with male harassment. This demonstrates how social niche construction can lead to higher level social structure; we suggest this approach could be used across a range of species in order to improve our understanding of the evolution of sociality.     

Cues for Mate Recognition and the Effect of Prior Experience on Mate Recognition in Enallagma Damselflies

In many coenagrionid damselflies, sexually mature females exhibit color polymorphisms, with some females resembling conspecific males. Although it has been suggested that the latter function as male mimics, this does not appear to be the case for those in the genus Enallagma. We found that sexually dimorphic coloration of the female abdomen and thorax are important cues for sexual recognition by males. We demonstrate for the first time in the Odonata, that males learn to recognize andromorphs as potential mates. After 2 days in an enclosure, sexually mature males exposed to only andromorphic females initiated more sexual interactions with tethered andromorphs than with heteromorphs, the majority morph in the natural population. Exposure to only heteromorphic females tended to decrease males' sexual responses to andromorphs, but not significantly so. Because the frequency of female morphs often varies within a population, learned mate recognition would be advantageous for males that search for mates. Our results lead to a novel, frequency-dependent hypothesis for the occurrence and maintenance of female-limited color polymorphisms.