Diurnal periodicity of spawning activity of permanently territorial damselfishes (Teleostei: Pomacentridae)

Synopsis Like other congeneric damselfishes, the herbivorous Stegastes altus defends individual feeding territories from heterospecific food competitors, regardless of sex. Females spawned demersal eggs for 31.4 min (n = 25) at the nest in territories of males 0–75 m away from theirs. Throughout the spawning season (June to September), spawning occurred only at dawn, mainly just after sunrise. Daily activity of the fish community showed that potential diurnal food competitors were few or inactive only at dawn and dusk. The frequency of intrusions into the female's territory by heterospecific competitors were as low when she left her territory to spawn at dawn, as they were in the daytime when she defended it. Removal of the female in the daytime resulted in a significant increase in intrusion frequency. An ‘anti-competitor hypothesis’, whereby dawn spawning ensures the food resource in the female's territory seems to explain not only the spawning periodicity in S. altus but also the timing of spawning of other permanently territorial damselfishes. Contribution from the Laboratory of Animal Ecology, Kyoto University, No. 492.     

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.     

Reproductive parasitism: male and female responses to conspecific and heterospecific intrusions at spawning in a mouth‐brooding cichlid Ophthalmotilapia ventralis

A rare form of alternative reproductive behaviour without simultaneous parasitic spawning was observed in Ophthalmotilapia ventralis, a lekking mouth‐brooding cichlid from Lake Tanganyika. Floater males attempted to sneak opportunistically into the territory to actively court the female, while the owner (bourgeois male) defended the territory against other potential intruders. Floater males had more body fat than territory owners and generally higher condition factors. In field experiments, the response of bourgeois males and courted females was tested towards floaters and egg predators (a catfish Synodontis multipunctatus) present in the territories. Territory owners responded aggressively particularly to floaters, and female responsiveness to bourgeois male courtship tended to decline when floaters were present. The potential influence of reproductive parasitism on sexual selection in mouth‐brooding cichlids is discussed.     

Ontogenetic changes in foraging tactics of the piscivorous cornetfish <Emphasis Type="Italic">Fistularia commersonii</Emphasis>

Foraging behaviors of the piscivorous cornetfish Fistularia commersonii were observed at shallow reefs in Kuchierabu-jima Island, southern Japan. This fish foraged on two types of prey fishes: one was reef fish that typically dwell on or near substrata (e.g., Tripterygiidae and Labridae), and the other was pelagic fish that shoal in the water column (e.g., Clupeidae and Carangidae). The prey sizes, prey types and foraging behaviors changed as the predator size increased. Prey sizes were largely limited by gape size of the cornetfish, and small predators consumed small prey. The small cornetfish (10–30 cm in total length) fed only on reef fish captured after stalking (where the fish slowly approaches the prey and then suddenly attacks). The stalking was done either solitarily or in foraging association with conspecifics. Large fish (30–120 cm) fed on both types of fishes by stalking and/or chasing (where the fish chases the prey using its high mobility and attacks), either solitarily or in foraging association with con- or heterospecifics. Thus, chasing was only performed by the large cornetfish against pelagic prey fish in associative foraging with other con- and heterospecific predators. As their body sizes increased, F. commersonii began to show a diversification of foraging behaviors, which was strongly related not only to the habitat types and anti-predatory behaviors of the prey fishes but also to associative foraging with con- or heterospecifics, which improves their foraging success.     

No evidence for learned mating discrimination in male Drosophila pseudoobscura

Background  

Since females often pay a higher cost for heterospecific matings, mate discrimination and species recognition are driven primarily by female choice. In contrast, frequent indiscriminate matings are hypothesized to maximize male fitness. However, recent studies show that previously indiscriminate males (e.g., Drosophila melanogaster and Poecilia reticulata) can learn to avoid heterospecific courtship. This ability of males to discriminate against heterospecific courtship may be advantageous in populations where two species co-occur if courtship or mating is costly.     

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.     

Spawning behavior and interspecific breeding in three japanese greenlings (hexagrammidae)

The spawing behavior ofHexagrammos otakii. H. octogrammus andH. agrammus was observed in two different regions of northern Japan using underwater video cameras placed near nests guarded by males. The spawning behavior of the three species consisted of similar patterns, although body size and nuptial coloration and nest location of territorial maleH. otakii differed from those of the other twoHexagrammos species. The courtship display of territorial males in each species involved rushing, butting and undulation of the trunk. When a female entered the nest, the male leaned his head on the future spawning bed in the nest and spasmodically undulated his trunk. The female that responded to the courtship laid her eggs within the seaweed bed. The territorial male then passed over the eggs, touching his genital pore to the egg mass, and released sperm. Sneaking by other males was frequently observed following the sperm emission. In both regions, females ofH. octogrammus andH. agrammus commonly responded to courtship of maleH. otakii and mated with them, but not vice versa. Possble reasons for the natural occurrence of such unidirectional hybridization are discussed.     

Brood defence and parental roles in a biparental cichlid fishLamprologus toae in Lake Tanganyika

Brood defence of a cichlid fish,Lamprologus toae, was investigated in its natural habitat in Lake Tanganyika. Both parents guarding a brood attacked both conspecific and heterospecific intruder fishes at different locations. The heterospecific intruder fishes could be classified into three groups on the basis of the locations at which the attacks against each species took place. The distinction of groups by the parents seemed to be primarily based on food habits and feeding behaviour of the intruder fishes. The piscivorous species which were more dangerous for the brood were attacked by both parents at more distant locations from the brood. Parental defence of breeding territory changed with the development of the young. The frequency of attacks against each group decreased after the young reached the size too large for the fishes of the group to prey on. Division of labour in the territorial defence was recognized between male and female. The male parent mainly defended the peripheral region of the territory and the female parent defended the inner region. Significance of the selective attack against intruders and the division of labour between the two sexes in brood defence is discussed.     

Alternative reproductive tactics in snail shell‐brooding cichlids diverge in energy reserve allocation

Life history theory predicts that the amount of resources allocated to reproduction should maximize an individual's lifetime reproductive success. So far, resource allocation in reproduction has been studied mainly in females. Intraspecific variation of endogenous energy storage and utilization patterns of males has received little attention, although these patterns may vary greatly between individuals pursuing alternative reproductive tactics (ARTs). ARTs are characterized by systematic variation of behavioral, physiological, and often morphological traits among same‐sex conspecifics. Some individuals may rely on previously accumulated reserves, because of limited foraging opportunities during reproduction. Others may be able to continue foraging during reproduction, thus relying on reserves to a lesser extent. We therefore predicted that, if male tactics involve such divergent limitations and trade‐offs within a species, ARTs should correspondingly differ in energy reserve allocation and utilization. To test this prediction, we studied short‐term and long‐term reserve storage patterns of males in the shell‐brooding cichlid Lamprologus callipterus. In this species, bourgeois males investing in territory defense, courtship, and guarding of broods coexist with two distinct parasitic male tactics: (1) opportunistic sneaker males attempting to fertilize eggs by releasing sperm into the shell opening when a female is spawning; and (2) specialized dwarf males attempting to enter the shell past the spawning female to fertilize eggs from inside the shell. Sneaker males differed from other male types by showing the highest amount of accumulated short‐term and long‐term fat stores, apparently anticipating their upcoming adoption of the nest male status. In contrast, nest males depleted previously accumulated energy reserves with increasing nest holding period, as they invest heavily into costly reproductive behaviors while not taking up any food. This conforms to a capital breeder strategy. Dwarf males did not accumulate long‐term fat stores at all, which they can afford due to their small behavioral effort during reproduction and their continued feeding activity, conforming to an income breeder strategy. Our data confirm that the resource storage patterns of males pursuing ARTs can diverge substantially, which adds to our understanding of the coexistence and maintenance of alternative reproductive patterns within species.     

A comparative study of territoriality in intertidal and subtidal blennioids (Teleostei, Blennioidei)

This paper presents a comparative analysis of territoriality in three intertidal (Lipophrys pholis, Coryphoblennius galerita, Salaria pavo) and two subtidal (Tripterygion delaisi, Parablennius pilicornis) blennioid fishes. Focal-animal observations of males guarding eggs showed that: (i) intertidal species had smaller territories that were less frequently patrolled; (ii) in subtidal species feeding was limited to the territory, while in intertidal fishes a substantial proportion of the feeding acts occurred outside the defended area; (iii) intertidal species spent less time out of the nest and showed a lower level of locomotory activity; (iv) subtidal species were subjected to a higher number of territorial intrusions with more species intruding their territory, some of them potential egg predators; (v) subtidal species performed several water column displays that played a significant role in courtship, while in intertidal fishes these displays are almost absent. Signalling without the presence of a female was almost absent in intertidal species. It is argued that intertidal fishes minimize the time spent out of the nest and the loss of contact with the substrate, features that may be of high survival value in conditions of strong turbulence. Although intertidal species have a reduced time available for feeding, this may be compensated by lower levels of locomotory activity, territorial defense and risk of egg predation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Social organization and spawning in the Atlantic sharpnose puffer,Canthigaster rostrata (Tetraodontidae)

Synopsis Social organization and spawning in the sharpnose pufferCanthigaster rostrataere studied on a reef in the San Blas Islands, Panama. Sexes were dimorphic. In mixed coral and rubble habitat, females defended territories against other females and small males. From one to six female territories were included within the territories of certain large males. These haremic males visited their females and patrolled their territories throughout the day. Smaller, non-haremic males occupied territories or home ranges within or adjacent to those of haremic males or were wanderers. Spawning between a haremic male and a territorial female occurred within the female's territory. The female prepared an algal nest into which demersal eggs were deposited. There was no parental care. Eggs were spherical, translucent, and measured approximately 0.66 mm in diameter. Larvae were about 1.4 mm TL and closely resembled those of other species ofCanthigaster.     

Territorial defense against various food competitors in the Tanganyikan benthophagous cichlid <Emphasis Type="Italic">Neolamprologus tetracanthus</Emphasis>

Territorial defense of nonbreeding female Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid, was investigated. Females defended territories (=home ranges, ca. 1m across) against a variety of intruding fishes. Conspecific females were usually attacked outside the territories, heterospecific benthivores (shrimp eaters) and omnivores near the border of the territories, and piscivores, algae and detritus feeders, and herbivores inside the territories. Females used some parts of the sandy substrate in the territories for foraging (foraging areas). Territorial defense prevented most of the conspecific females and benthivores from intruding into the foraging areas. In omnivores, piscivores, and algae and detritus feeders, about half the intruders were repelled from the foraging areas, although herbivores were infrequently repelled in the areas. Soon after removal of the resident females, many food competitors invaded the foraging areas and eagerly devoured prey, suggesting that the territories are maintained for food resource protection from these competitors. Females are likely to discriminate intruding fishes and change their territorial defense primarily on the basis of the degree of dietary overlap, resulting in monofunctional serial territories.     

Reproductive interference by male Drosophila subobscura on female D. persimilis: A laboratory experiment

While females often reject courtship attempts by heterospecific males, reproductive interference by harassment from such males can nonetheless occur, potentially reducing female fitness. Such effects may be profound following a range expansion, when males from a new species may suddenly encounter (and perhaps even become abundant relative to) females of related native species. Drosophila subobscura recently invaded North America and may impact native species through reproductive interference and other processes. We test for the potential for reproductive interference by D. subobscura males on D. persimilis females in the laboratory. D. subobscura males aggressively copulated with D. persimilis females, including many females that exhibit rejection behaviors. Despite females attempting to dismount the males, the heterospecific copulations are on average longer than conspecific copulations, and females exhibit some reluctance to remate with conspecific males following this harassment. Females confined with both conspecific and heterospecific males produce fewer adult progeny than those with either conspecific males only or with conspecific males and distantly related D. simulans males that do not engage in female harassment. Overall, our results illustrate how reproductive interference by an invasive species can have negative effects on resident natural populations.     

Nest use by territorial males in a shell-brooding cichlid: the effect of reproductive parasitism

Male Telmatochromis vittatus, a substrate-brooding Tanganyikan cichlid, exhibit two parasitic reproductive tactics: takeover of spawning by larger males (pirates) and sneaking by smaller males (sneakers). Medium-sized males are territorial and pair-spawn within nests of clumped shells that harbor several resident females that are potential mates of the territory owner. To study nest use by territorial males, we analyzed the relationship between the body size of territorial males and nest quality, the number of females per nest, the distance between nests, and the frequency and intensity of reproductive parasitism. The size of males was not correlated with nest quality, the number of females, or the number of sneakers, but was negatively correlated with the frequency of intrusion by pirates and was positively correlated with the distance between nests. Territorial males effectively defended nests against sneakers but failed to defend against pirates. These results suggest that larger territorial males selected nests that have a lower risk of usurpation of spawning. We hypothesize that the risk of intrusion by pirate males affects the selection of nests by territorial males in this species.     

Spawning ecology of the American brook lamprey, <Emphasis Type="Italic">Lampetra appendix</Emphasis>

Synopsis We observed spawning American brook lamprey, Lampetra appendix, in coldwater streams in Minnesota to assess various aspects of their spawning behavior and spawning habitat requirements. Spawning occurred during April and May, at water temperatures ranging from 8.7 to 15.5°C. Average adult lamprey length and mass differed significantly among streams, but there were no significant differences in length or mass between males and females. Overall sex ratio was 1:1, although one stream had significantly more males than females and one stream held significantly more females. Lampreys spawned in groups of 2–14 individuals, averaging 4.2 adults per nest across all streams. Nests were constructed in gravel and cobble substrate just upstream of riffles, spaced at an average density of three nests m⁻². The typical nest was 16 cm in diameter in water 31 cm deep with a bottom current velocity of 14 cm s⁻¹, and was excavated to a depth of 4 cm below the stream bottom; however, some nest characteristics varied significantly in a few streams. Nests were larger in streams with larger spawning groups, deeper water, and slower current velocities. American brook lamprey exhibited spawning behaviors and spawning habitat requirements similar to those of other species of lamprey in North America.     

Sex-role reversal in the tidewater goby, Eucyclogobius newberryi

Species in which females compete more intensely than males for access to mates are uncommon. Sex-role reversal in fishes has been documented only in species in which males bear eggs, such as pipefish and a mouth brooding cardinalfish. I investigated the reproductive behavior of the tidewater goby, Eucyclogobius newberryi (Gobiidae), to determine whether and to what degree this species is sex-role reversed. Males constructed and defended burrows for spawning in sand. Both sexes initiated courtship, but the female's breeding coloration was more striking. The intensity of sexual aggression was greater among females than among males. The female laid her entire clutch with a single male, and the male accepted only one clutch per brooding cycle. Both sexes spawned repeatedly (up to 12 times in aquaria), but fish did not form permanent pairs. Males cared for eggs in the burrow 9–11 days until hatching, and rarely if ever emerged to feed. Many aspects of male behavior (nest construction and defense, courtship, and parental care) were typical of most gobiids. On the other hand, female behavior (black nuptial coloration and intense female-female competition) was unusual, not only for gobiids but for animals in general. I therefore concluded that the tidewater goby is moderately sex-role reversed. Its sexual behavior is apparently unique among fishes because it is the only reported case of sex-role reversal in teleost males that do not bear eggs or developing young. This revised version was published online in July 2006 with corrections to the Cover Date.     

Breeding ecology and nest aggregations in a population of Salaria pavo (Pisces: Blenniidae) in an area where nest sites are very scarce

In this paper we present data on the breeding ecology and patterns of nest aggregation in a population of Salaria pavo (Pisces: Blenniidae), in an area where hard substrates are very scarce. The study site was at Ria Formosa, an extensive littoral lagoon on the south coast of Portugal. The only hard materials available are bricks, tiles, stones and debris that clam culturists use to delimit their fields. The males of S. pavo establish nests intertidally in the holes of the bricks where they guard the eggs. Data were collected at low tide inspections of these artificial ridges and underwater behavioural observations during high tide. The main results are: (i) the same male can establish sequentially more than one nest in the same breeding season; the number of males that stay within the bricks increases before the start of the breeding season and declines towards its end; (ii) there is a large excess of mature males that do not establish nests and they are significantly smaller than the nesting males, suggesting very strong competition for nest sites; (iii) the nests of several males occur in adjacent holes of the same brick, sometimes being entirely surrounded by other nests. Thus, in this population, there is virtually no defended territory around the nest, a situation not known for other blenniid species. This pattern of nest aggregation implies both strong competition among males and reduction of aggression between neighbour parental fishes. This can be explained by the extreme scarcity and spatial distribution of available nest sites.     

Variation of territory size and defense behavior in breeding pairs of the endemic Lake Tanganyika cichlid fish Variabilichromis moorii

Lake Tanganyika harbors the ecologically, morphologically, and behaviorally most diverse species flock of cichlid fishes. It is comprised by substrate breeding and mouthbrooding species, most of which live in littoral habitats. Species communities are characterized by complex behavioral and trophic interactions, resulting in a dense pattern of partially overlapping territories, depending on the degree of ecological distinctness. We studied territorial behavior of breeding pairs in a substrate breeding species, with respect to territory size and defense behavior. The study species Variabilichromis moorii belongs to the tribe Lamprologini, the most species rich tribe of cichlids in Lake Tanganyika. Our study shows that breeding pairs of V. moorii can have highly complex territories, in which both parents hold separate sub-territories which are shifted slightly according to the movements of the fry, but the outer borders are conjointly defended. The size of the total defended territory varied from <1 to almost 4 m², averaging at about 2 m². Depending on presence of competitors or fry-predators evoking agonistic interactions, the territory size varied quite substantially over the day. Attack rates and size of the defended area decreased with water depth. Agonistic behavior was observed toward heterospecifics as well as conspecifics, with heterospecific attacks mostly concerning territorial neighbors and potential fry-predators in about equal frequencies. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: T. Wilke, R. V?in?l? & F. Riedel Patterns and Processes of Speciation in Ancient Lakes: Proceedings of the Fourth Symposium on Speciation in Ancient Lakes, Berlin, Germany, September 4–8, 2006     

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.     

Competition for spawning sites between two gobiid fishes, Bathygobius fuscus and Eviota abax, derived by alteration of mating tactics of the former

Interspecific competition for spawning sites between two gobiid fishes, Bathygobius fuscus and Eviota abax, was studied on a rocky shore. Large males (LM: 55–80mm in standard length: SL) and small males (SM: 34–52mm SL) of B. fuscus acted as nest holders and sneakers, respectively, in the early spawning season (July). The sympatric male E. abax (22–33mm SL) was smaller than SM B. fuscus, and synchronically used rocky holes much smaller than those of LM B. fuscus. In this season, aggressions between the two species were rarely observed. In late season (August), as the number of the LM decreased, the SM converted their tactics to nest holding, occupying nests of a sizes similar to those of E. abax. Eviota abax males were dispossessed by SM and shifted their nest sites to cavities between cobbles and sandy bottom, which might be poorer nest sites than rocky holes.