Alternative reproductive tactics and courtship in the common goby

In the marine common goby Pomatoschistus microps , males either build nests and attract females to spawn or act as sneakers, forcing themselves into other males' nests during mating, to fertilize some of the eggs in the clutch. The reproductive tactics of small males were studied in the presence of surplus nest material and an excess of gravid females, while a larger, nestholding male spawned with a female. Of 24 small males, 13 only attempted to sneak, seven only courted females and/or spawned in their own nest, and four performed both behaviours. The small males who attracted females successfully were significantly larger than the ones who failed to do so. Females had been courting the small male more often when he subsequently spawned in his own nest, than when he did not get a female. The small males did not court the females very frequently, and their courtship rate did not depend on whether or not they attracted females. Instead, female courtship behaviour may be a cue for a male either to spawn as a nest holder or become a sneaker. The study pointed out the importance of female behaviour for male reproductive tactics, which could be more important in fish than is known currently.     

Male body size and brooding success under natural conditions in the freshwater Japanese goby, Rhinogobius sp. OR (orange type)

Males of the Japanese goby Rhinogobius sp. OR (orange type) establish nests under stones in streams. They provide care to egg masses until hatching. We carried out field observations and a field experiment in a natural stream to examine the effect of male body size on brooding success. An experiment excluding guarding males indicated that their presence has a positive effect on nest defense. Field data showed that male body size was positively correlated with the size of the egg mass that was initially found at the nest. Furthermore, a sequential increase in egg mass size seen in the nests of larger males indicated that multiple females spawned during a single breeding cycle of larger males. To examine male brooding success, two dependent variables, egg mass persistence (days from initial egg mass to disappearance or hatching) and hatching success were analyzed. The results indicated that the variation found in both dependent variables was significantly explained by two independent variables, namely, initial egg-mass size and the number of matings. It is also suggested that male body size indirectly influenced brooding success through positive correlations with these independent variables.     

Nest Building and Mating in Relation to Male Size in Corkwing Wrasse, Symphodus Melops

This study examines nest building and mating in relation to size in male corkwing wrasse, Symphodus melops, in the absence of male–male competition. We kept males of varying size together with mature females in tanks containing adequate materials and sites for nest construction. Our results indicate that the size of territorial males does not influence the ability to construct a nest. However, larger males initiated nest building earlier than smaller males and nest size correlated positively with male size. Furthermore, male size did not affect whether or not a nesting male succeeded in mating, indicating that male size was not essential for female acceptance (i.e. whether or not a female is willing to mate) of a male in the current situation.     

Male–male competition for large nests as a determinant of male mating success in a Japanese stream goby, Rhinogobius sp. DA

Males of the stream goby Rhinogobius sp. DA (dark type) court females in deep pools and care for the eggs under stones in shallow riffles. We studied male–male competition for access to females and nest sites to understand how male size influences the mating success of this species. In field observations, larger males won in fighting with other males. However, large males did not tend to monopolize courtship opportunities, and the frequency of successful courtships, after which males led the females to the nests, was not related to male body size. The fact that courted females always escaped from the fighting sites once males began fighting likely explains why male size was not positively related to courtship success. Large males occupied large nest stones, and the number of eggs received in the nest was correlated positively with nest size. In aquarium experiments with two tiles of different sizes provided as nesting materials, males always chose the larger nest and, when two males were introduced simultaneously, the larger one occupied the larger nest. These results suggested that male mating success of this goby is determined by male–male competition for large nests rather than for access to females. Received: June 9, 2000 / Revised: September 2, 2000 / Accepted: October 4, 2000     

The Race for Females: The Mating System of the Red Mason Bee, Osmia rufa (L.) (Hymenoptera: Megachilidae)

The ecology of a species strongly influences the strategies with which males and females maximize their lifetime reproductive success. When males and females do not invest equally in offspring, the sex with the higher parental investment becomes a rare resource for the other. The spatial and temporal distribution of the limiting sex forms the basis of the mating system. In nest-constructing Aculeata such as the red mason bee, Osmia rufa, females perform intensive brood care, whereas males do not invest in their offspring but instead compete for access to mates. Receptive females of this species are widely distributed and do not assemble at certain places. Therefore, territorial behavior is not an advantageous mating tactic for males, which instead search for females within individual home ranges usually centered around food plants. The unmarked or defended home ranges of different males may completely overlap. Competitive searching leads to a random distribution of matings among males that is largely independent of body size. The mating system of O. rufa can be described as scramble competition polygyny.     

Size-mediated dominance and aggressive behavior of male Japanese fluvial sculpin Cottus pollux (Pisces: Cottidae) reduce nest-site abundance and mating success of conspecific rivals

Despite growing evidence for plasticity in the mating patterns of nest-holding animals in relation to the changes in nest abundance, the effects of aggressive interaction by dominant males on nest availability for conspecific rivals remains unclear. To quantify the effects of male–male competition on nest-site choice and mating success of the male Japanese fluvial sculpin Cottus pollux, we conducted experiments on 5 males from different 5 size classes under both sufficient and shortage nest-abundance conditions. Nest-choice experiments showed that both male size class and nest-abundance condition had significant effects on the nesting rates of males. Following the nest-choice experiments, 10 gravid females were added in the experimental tanks. Mating experiments revealed that male size, nesting rate before addition of females, and the number of courtship attempts on females were valid variables of male mating success, regardless of nest-abundance conditions. After achieving initial mating success, the largest nesting male exhibited more frequent aggressive interaction with other conspecific males than he did before obtaining eggs in his nest. Our results suggest that size-mediated dominance and aggressive behavior of males may disrupt nest acquisition of other conspecific males, and may consequently result in extreme variation in mating success among males even under sufficient nest-abundance conditions.     

Mate availability affects female choice in a fish with paternal care: female counterstrategies against male filial cannibalism

Theory predicts that individuals should adopt counterstrategies against intersexual conflict with their mating partners if the counterstrategies are effective and cost-efficient. In fishes, males with parental care often cannibalize their own offspring, which reduces the female’s fitness and creates intersexual conflicts. Males of the goby Rhinogobius flumineus cannibalize more eggs in the nest when they have access to additional females prior to spawning. Thus, it is predicted that females will strategically avoid spawning with males that have high mate availability. In the present study, we experimentally tested this prediction. When sexual pairs were placed in tanks, most females (control females; 21/22) successfully spawned inside the nest. In contrast, when a gravid female (stimulus female) that was housed in a small transparent cage was shown to the experiment pairs prior to spawning, only about half of the females (experiment females; 16/29) spawned inside the nest; the remaining females released unfertilized eggs outside of the nest. Moreover, experiment females infrequently accepted and followed males into nests, and delayed spawning more often than control females. R. flumineus females prefer males that court frequently. Indeed, experiment females that infrequently received courtship tended to spawn outside of the nest. However, infrequent courtship alone could not explain outside-nest spawning, delay in spawning, or the shorter stay of females in nests. These results imply that the presence of a stimulus female dampens female spawning with males. We suggest that R. flumineus females may strategically reject or hesitate to spawn with males that have high mate availability, and that this spawning avoidance may be a counterstrategy against male filial cannibalism.     

Size and overlap of home range inCalomys musculinus (Muridae: Sigmodontinae)

Home range size and overlap ofCalomys musculinus (Thomas, 1913) was examined in relation to sex and breeding periods. The study was carried out in four 0.25 ha enclosures, in a natural pasture, between October 2002 and July 2003. The four enclosures functioned as independent populations and each was considered a replicate. The capture, mark and recapture method was used. Home range size and overlap inC. musculinus depended on sex and period. Home range size of males was always larger than that of females. Females showed a small degree of intra-sexual home range overlap during the breeding period. In general, male home ranges overlapped with females. We conclude that differences in home range size inC. musculinus is determined by sex and breeding period. Moreover, the degree of inter- and intra-sexual home range overlap during the breeding period suggests that both males and females ofC. musculinus use space differently. Females did not share their home range with other females, while males fully shared it with both sexes, and male spacing is influenced by the distribution of females. A promiscuous-polygynous mating system is suggested forC. musculinus.     

Spacing Behavior of a Non‐Larder‐Hoarding Tamiasciurus: A Study of Mearns's Squirrels in Xeric Coniferous Forests

In ecosystems with seasonal fluctuations in food supply many species use two strategies to store food: larder hoarding and scatter hoarding. However, because species at different geographic locations may experience distinct environmental conditions, differences in hoarding behavior may occur. Tree squirrels in the genus Tamiasciurus display variation in hoarding behavior. Whereas red (Tamiasciurus hudsonicus) and Douglas's (Tamiasciurus douglasii) squirrels in mesic coniferous forests defend territories centered around larder hoards maintaining non‐overlapping home ranges, red squirrels in deciduous forests defend small scatter‐hoarded caches of cones maintaining overlapping home ranges. As in other rodent species, variation in hoarding behavior appears to influence the spacing behavior of red and Douglas's squirrels. In contrast, Mearns's squirrels (Tamiasciurus mearnsi) in xeric coniferous forests neither rely on larder hoards nor appear to display territorial behavior. Unfortunately, little is known about the ecology of this southernmost Tamiasciurus. Using radiotelemetry, we estimated home‐range size, overlap, and maximum distance traveled from nest to examine the spacing behavior of Mearns's squirrels. Similar to scatter‐hoarding rodents, maximum distance traveled from nest was greater for males during mating season, whereas those of females were similar year round. Although no seasonal differences were detected, male home ranges were three times larger during mating season, whereas those of females were smaller and displayed a minor variation between seasons. Home ranges were overlapped year round but contrary to our expectations, overlap was greater during mating season for both sexes, with no detectable relationship between male home‐range size and the number of females overlapped during mating season. Overall, the results appear to support our hypothesis that in the absence of larder hoards, the spacing behavior of Mearns's squirrels should be different from larder‐hoarding congeners and more similar to scatter‐hoarding rodents.     

Influence of male mating history on female reproductive success among monandrous Naryciinae (Lepidoptera: Psychidae)

1. Multiple male copulations can have detrimental effects on female fitness due to sperm limitation. 2. Monandrous Naryciinae females are immobile while the males are short‐lived and do not feed. Multiple male mating is therefore expected to lead to sperm limitation in females. Sperm limitation and male limitation are hypothesised as causes of the repeated evolution of parthenogenetic reproduction in the Psychidae. 3. In this study, the effects of multiple male mating on female reproduction are investigated in several species of Naryciinae by allowing males multiple copulations. The results for two species, Siederia listerella and Dahlica lichenella, are compared. The sex ratios of 53 natural populations are examined for indications of male limitation. 4. Previous copulations by the male increased the female's risk of remaining unfertilised. However, contrary to expectations, those unfertilised females were capable of successful re‐mating. 5. In S. listerella, the number of previous copulations of males negatively influenced female fitness. Females produced 30% fewer offspring if they mated with a previously mated male. In D. lichenella, the older the male and the lower its number of total lifetime copulations, the higher the female's reproductive success. 6. Only a fraction of the investigated populations had a female‐skewed sex ratio, but differences in development time between males and females could lead to reproductive asynchrony. 7. In conclusion, male mating history did not lead to strong sperm limitation in Naryciinae as had been suggested by their life history.     

Timing of Spawning and Host-nest Choice for Brood Parasitism by the Japanese Minnow,Pungtungia herzi,on the Japanese Aucha Perch,Siniperca kawamebari

A field study of the breeding ecology of the Japanese aucha perch, Siniperca kawamebari, and brood parasitism by the Japanese minnow, Pungtungia herzi, on nests of the perch was carried out from 1989 to 1991. Observations of perch nests under natural conditions in 1990 showed that brood parasitism by the minnow was concentrated on host nests in which nest owners had just begun their nesting cycle. When spawned in a perch nest with recently spawned perch eggs, parasite eggs always hatched earlier than host eggs. An experiment with imitation perch eggs in 1991 confirmed that changing colour of host eggs was the cue for the parasites to distinguish between different developmental stages of host eggs. Parasite eggs rapidly disappeared without guarding by a host male (Baba et al. 1990). This loss was caused by predation by fishes. Parasite fry left the nest immediately after hatching, so parasite eggs spawned in a host nest in an early stage should be well guarded until they hatch. In the field, minnows deposited their eggs in perch nests which had larger numbers of newly spawned perch eggs. Since the perch males always deserted their nests when their own eggs disappeared, the parasite's choice of host nests with larger numbers of host eggs may ensure survival of the parasite eggs. The timing of egg deposition and choice of host nest by the minnow appear to be adaptive in terms of brood parasitism on nests of the perch.     

Spacing pattern in a social group of stray cats: effects on male reproductive success

Reproductive consequences of male spacing patterns have received relatively little attention in nonterritorial mammals, in particular in group-living species, where most studies have focused on the relation between social rank and reproductive success. We investigated the effects of spacing pattern on male reproductive success within a social, nonterritorial, promiscuous population of stray cats, Felis catus. Male home ranges overlapped home ranges of many females, consistent with a promiscuous mating system. Furthermore, males with the largest home ranges included the most female home ranges; they successfully reproduced with these females and had the highest reproductive success. Home range size predicted male reproductive success even when controlling for the effect of social rank. However, males also reproduced with females whose home range did not overlap their home range, suggesting that males can make quick excursions outside their home range to find new mating opportunities. We conclude that, in group-living situations, a male's ability to maintain a large home range may be one of the principal causes of variation in mating success in the stray cat.     

Simultaneous spawning by female stream goby Rhinogobius sp. and the association with brood cannibalism by nesting males

A laboratory experiment was conducted by varying the undersurface area of nesting substratum and the number of females in an experimental tank to elucidate the determinants of the mating pattern in the stream goby, Rhinogobius sp. cross‐band type. Males with larger nests tended to attract two or more females to their nest in a tank. Moreover, males spawned simultaneously with multiple females and entire brood cannibalism by males was rarely observed under a female‐biased sex ratio. When males spawned with a single female with low fecundity, however, entire brood cannibalism occurred at a high frequency, suggesting that a male guarding a nest with fewer eggs consumes the brood. Therefore, spawning behaviour of females that leads to a large egg mass would decrease the risk of entire brood cannibalism. In this species, simultaneous spawning by multiple females in a nest serves as a female counter‐measure against entire brood cannibalism. These results suggest that a conflict of interest between the sexes through brood cannibalism is a major determinant of simultaneous spawning.     

Affiliative relations among male Japanese macaques (Macaca fuscata yakui) within and outside a troop on Yakushima Island

Male Japanese macaques (Macaca fuscata yakui) in a troop on Yakushima Island frequently groom other males. However, previous studies have not compared the social relations of troop males to those of non-troop males. I followed all troop males and non-troop males in and near a troop during a mating season and during the following non-mating season and recorded their neighbors, grooming, and agonistic interactions. Comparisons of the social relations of troop males and non-troop males with other troop members revealed that grooming and agonistic interactions with females during the mating season were similar between troop and non-troop males. However, troop males groomed each other more often and had fewer agonistic interactions among themselves than did non-troop males. Compared to what occurred in the mating season, troop males groomed females less often and exchanged grooming bouts more often with other troop males during the non-mating season. One non-troop male groomed females more frequently than did any troop male in both seasons, and this male groomed troop males more frequently than did any troop male in the non-mating season. This male immigrated into the troop during the following mating season. Regardless of their competition with respect to reproduction, male Japanese macaques on Yakushima Island maintain affiliative relations, probably to cooperatively defend fertile females from non-troop males.     

Male mating success and female mate choice in the river sculpin,Cottus nozawae (Cottidae)

Synopsis Mating success of males and its correlates were investigated in a natural population of the polygynous fluvial sculpinCottus nozawae. Furthermore, the female mate preference of this species was examined experimentally under alternative conditions for mating in a stream. The mating success of individual males (the number of females with which a male mated) ranged between 0 and 8 with a mean of 2.41 in 1983 and 2.52 in 1989, in a population of which the sex ratio was about 1 : 2 in both years, skewed toward females. Mainly due to the excess of nests without egg masses and the few nests with one egg mass, the distribution of male mating success did not fit a Poisson distribution, indicating its non-randomness. Male mating success was not correlated either with the size of the nest rocks or with the male size, suggesting that these two variables are not determinants of mating success. The mate choice experiments demonstrated that females of this species more frequently chose smaller males as mates whose nests already contained eggs than large males without eggs. Additionally, an analysis of stomach contents of guarding males suggested that the parental males ate their own eggs during egg guarding (filial-cannibalism). Based on these results and on a comparison of reproductive characteristics with congeneric species, it is suggested that one of the most important determinants for female mate choice inCottus species may be whether or not parental males are filial egg cannibals.     

Female Mate Preference and Male Mating Success of Ayu Fish,Plecoglossus altivelis (Osmeridae) under a Promiscuous Mating System

The ayu, Plecoglossus altivelis, is known to show wide variation in adult body size. We examined the spawning behavior in experimental spawning groups of fish in which male body size varied. Males never competed with each other for females or spawning site, while females spawned repeatedly, 17–97 times depending on body size, with previous or novel males. Males exhausted their sperm after a single mating episode, on average, regardless of body size. Moreover, repeated sperm production apparently reduced the residual lifespan. Females preferred mating simultaneously with more than one male and allowed males of body size similar to their own to mate more frequently. Thus the largest male within a spawning group was not always the most successful at mating, but mating success of any given male appears to depend upon body size distribution of females within the population. Female mate preference has apparently evolved to ensure complete fertilization under circumstances where males have been selected to economize sperm output during any one mating episode.     

Monogamous Mating System and Spawning Cycle in the Gobiid Fish, Amblygobius phalaena (Gobiidae)

Reproductive ecology and mating system of the gobiid fish, Amblygobius phalaena, were studied on the coral reef at Sesoko Island, Okinawa, Japan. This goby usually lives in pairs, and maintains territories with several burrows for shelter and spawning. Although a few paired individuals changed partners, most pairs remained together over successive rounds of reproduction. Mate guarding by females appeared to prevent males from mating with other females. Spawnings were synchronous with semilunar periods. Several expected spawnings failed to occur (12%). These may have been caused by the delays in spawning preparation of the paired females or by the disturbance caused by a typhoon. A pair spawned in one of the several burrows within their home ranges. Eggs were deposited on the ceiling of the burrow, and were tended by the paired male for 3–4 days until embryos hatched. The males tended eggs at the expense of their feeding. Aggression toward fishes approaching their burrows were exhibited by the males as well as the females. Because of its low frequency in females, this behavior did not limit their ability to feed.     

Male mating success in the Azorean rock-pool blenny: the effects of body size, male behaviour and nest characteristics

Larger male Azorean rock-pool blennies Parablennius sanguinolentus parvicornis received more female visits, were courted more often by females and received more spawnings. Larger males also received a higher number of male intrusions, attacked more conspecifics and defended larger territories. Larger males showed more nest cleaning behaviour and a marginally non-significant trend for higher egg fanning rate. Male courtship, male attack rate against conspecifics and parental behaviour were all correlated with the frequency of female spawnings received by each male even when controlling for male size, suggesting that these behaviour patterns influence male mating success. On the other hand, a positive partial correlation was found between female courtship and the frequency of female spawnings, controlling for the number of female visits, which suggests a role for female mate choice on male mating success. Finally, males nesting in chambers in the bottom of pools received more spawnings than males nesting either in crevices or under boulders. However, nest opening area was associated significantly negatively with male mating success, when controlling for male size. Thus, the present data suggest strongly that male characteristics overrule nest characteristics in determining male mating success in the Azorean rock-pool blenny.     

Polygamous mating system and protogynous sex change in the gobiid fish <Emphasis Type="Italic">Fusigobius neophytus</Emphasis>

Whereas mating behaviors and social structure have been studied extensively in monogamous hermaphroditic gobiid species, such studies are relatively limited for polygamous gobiid species. To investigate the reproductive strategy of polygamous gobies, mating groups of the common fusegoby Fusigobius neophytus were observed on reefs of Kuchierabu-jima Island, southern Japan. Males established mating nests on flat-rock surfaces within their territorial home ranges on sandy rubble flats. Females maintained independent home ranges outside the male home ranges during nonreproductive periods, but they shifted their home ranges to overlap with male ranges and actively visited male mating nests during their reproductive periods (1–3 days at ca. 7-day intervals). Females often changed mates during their serial mating. The mating system used by the common fusegoby fits with the definition of male-territory-visiting polygamy. The sex ratio within the study population was female-biased. Nest-holding males were significantly larger than females and were polygynous (mating with up to eight females). These characteristics fit well with the prediction of protogyny by the size-advantage model. Some of the females were observed to undergo functional sex changes to nest-holding males. In addition, small floating males demonstrated sneaking behavior. None of the floating males were derived from females that had changed sex, suggesting a diandric life-history pathway for F. neophytus.     

Alternative mating tactics of the gobiid fish <Emphasis Type="Italic">Bathygobius</Emphasis><Emphasis Type="Italic">fuscus</Emphasis>

The reproductive ecology of the gobiid fish Bathygobius fuscus was studied at Nobeoka, Miyazaki, Japan. Males of this species maintain small rock holes as a nest and females spawn an egg mass on the wall of the nest. The males employed two forms of mating tactic: nest holding and sneaking. A nest holder stayed in the nest and waited for a female to visit, whereas a sneaker intruded into a nest while a pair was engaged in reproduction. Males larger than 55 mm standard length were always nest holders; those of smaller size employed both tactics. As the larger males excluded the smaller males, the latter did not occupy a nest hole. With a decrease in the number of larger males, smaller males changed their mating tactic from sneaking to nest holding. The results suggest that male Bathygobius fuscus adopt a conditional strategy whereby they change their tactic depending on their social status. Electronic Publication