Size-dependent male mating behaviour and sexual harassment in a population of Atlantic mollies (Poecilia mexicana) from a sulphur creek

The Atlantic molly Poecilia mexicana inhabits a variety of habitats. In previous studies, small males from clear-water populations showed considerably more sexual behaviours than large males. Males from a sulphur creek, the El Azufre, or from a sulphur cave (cave molly) showed comparatively low sexual activity, and typical size-dependent mating behaviour was either absent or large males showed even more sexual behaviour than smaller ones. Sexually deprived cave molly males increased their sexual activity, but small males still did not show more sexual behaviour than large ones, suggesting that the genetic basis for typical size-dependent mating behaviour has been lost. Likely, the extreme abiotic conditions (hypoxia and toxic hydrogen sulphide) have selected against energetically costly behaviours. In this paper, we examined how El Azufre males would respond to sexual deprivation. We found these males to also increase their sexual activity after sexual deprivation but, unlike cave mollies, small males now showed more sexual behaviour than large ones, suggesting that the genetic basis for this behaviour is not lost in this population. Differences between populations may be due to the less harsh characteristics of El Azufre (e.g. less H₂S), resulting in weaker selection against energy-consuming behaviours. Furthermore, we tested for potential costs of male sexual harassment for females in terms of reduced feeding efficiency in the presence of a male, as was shown for P. mexicana from a clear-water population, but was not found in cave mollies or the El Azufre fish. Test females were either starved for 1 week to increase hunger levels or male sexual deprivation and female starvation were combined; however, we found no evidence for male sexual harassment even after these treatments. Our results parallel previous findings in the cave molly and suggest that El Azufre males have lost specific behavioural traits that cause sexual harassment.     

Sexual harassment in a live-bearing fish (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>): influence of population-specific male mating behaviour

We investigated male sexual behaviour and the cost of sexual harassment, as measured by the reduction of female feeding time in the presence of a male, in a cave-dwelling population of Poecilia mexicana, in which sexual harassment does not occur naturally. We asked whether the lack of sexual harassment in this population is due to low sexual activity of the males, or low feeding motivation of the females. We experimentally increased the sexual activity of males or the females feeding motivation, or we used a combination of both treatments. Female feeding time was not lower in the presence of a male than in the presence of a female after sexual deprivation of the males or food deprivation of the females. Only in the combined experiment was female feeding time lower in the presence of a large male than in the presence of a small male, indicating a weak effect of sexual harassment by large males. Virgin females did not suffer a cost of sexual harassment, indicating that sexual experience does not cause the lack of sexual harassment in cave mollies. Males from a surface population, where sexual harassment occurs, significantly reduced the feeding time of cave-dwelling females even though these males exhibited surprisingly little sexual behaviour. The sexual activity of cave mollies did not correlate with male body size in any experiment, indicating that even after sexual deprivation, small cave molly males do not switch to the alternative mating behaviour known in surface-dwelling P. mexicana, where sexual activity is correlated negatively with male body size.     

Predation of a cave fish (Poecilia mexicana, Poeciliidae) by a giant water-bug (Belostoma, Belostomatidae) in a Mexican sulphur cave

Abstract.  1. Caves are often assumed to be predator-free environments for cave fishes. This has been proposed to be a potential benefit of colonising these otherwise harsh environments. In order to test this hypothesis, the predator–prey interaction of a belostomatid (predator) and a cave fish (prey) occurring in the Cueva del Azufre (Tabasco, Mexico) was investigated with two separate experiments.
2. In one experiment, individual Belostoma were given a chance to prey on a cave fish, the cave form of the Atlantic molly ( Poecilia mexicana ), to estimate feeding rates and size-specific prey preferences of the predator. In the other experiment, population density of Belostoma was estimated using a mark–recapture analysis in one of the cave chambers.
3. Belostomatids were found to heavily prey on cave mollies and to exhibit a prey preference for large fish. The mark–recapture analysis revealed a high population density of the heteropterans in the cave.
4. The absence of predators in caves is not a general habitat feature for cave fishes. None the less predation regimes differ strikingly between epigean and hypogean habitats. The prey preference of Belostoma indicates that cave-dwelling P. mexicana experience size-specific predation pressure comparable with surface populations, which may have implications for life-history evolution in this cave fish.     

Sex recognition in surface- and cave-dwelling Atlantic molly females (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, Poeciliidae,Teleostei): influence of visual and non-visual cues

Cave fishes need to rely on non-visual senses, such as the sense of smell or the lateral line to communicate in darkness. In the present study, we investigated sex identification by females of a cave-dwelling livebearing fish, Poecilia mexicana (cave molly), as well as its surface-dwelling relatives. Unlike many other cave fishes, cave mollies still possess functional eyes. Three different modes of presentation of the stimulus fish (a male and an equally sized female) were used: (i) the stimulus fish were presented behind wire-mesh in light, allowing the focal female to perceive multiple cues, (ii) the experiment was carried out under infrared conditions, such that only non-visual cues could be perceived and (iii) the stimulus fish were presented in light behind transparent Plexiglas, allowing for the use of visual cues only. Females of all populations examined preferred to associate with the stimulus female in at least one of the treatments, but only when visible light was provided, suggesting that far-range sex recognition is limited or even absent in the cave molly under naturally dark conditions.     

Sperm production in an extremophile fish,the cave molly (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, Poeciliidae,Teleostei)

A prominent trade-off in life history theory and evolution balances the costs of reproduction with those of basic somatic needs. Hence, reproductive efforts may be reduced in environments where additional energy is required for somatic maintenance. Here, we investigated male sperm stores in Atlantic mollies (Poecilia mexicana) from a sulfidic cave and several sulfidic and non-sulfidic surface habitats. We found significant differences among populations in the number of sperm stripped per male, which was also correlated with differences in gonad weights. The largest sperm stores were detected in males from non-sulfidic surface creeks, while males from a partially sulfidic surface system had lower sperm counts, and males from completely sulfidic systems, surface as well as subterranean, had even fewer available sperm. We conclude that the extreme environmental conditions in sulfidic habitats appear to constrain male sperm production, since hydrogen sulfide as a naturally occurring toxin requires energy-demanding adaptations. Furthermore, we examined sperm counts of lab-reared cave and surface mollies in response to energy limitation. Males from stock populations were placed under high and low food treatments for a 2-week period and then stripped of sperm. Sperm counts of surface mollies tended to be reduced by low food availability, whereas sperm counts of cave mollies did not significantly vary between food treatments, which likely points towards a higher starvation resistance in cave mollies.     

Size and morphological variation in males of the sailfin molly,Poecilia latipinna

Synopsis Mature males of the sailfin molly, Poecilia latipinna (Poeciliidae), display extensive variation in size and morphology. Small males, less than about 30 mm SL, usually exhibit no distinctive color or morphological development. Large males, greater than about 45 mm SL, display full expression of the sailfin traits, including bright blue and gold life colors, distinctive pigment markings on the fins and body, and a greatly enlarged dorsal fin. Intermediate-sized males, 30–45 mm SL, display intermediate and variable development of secondary sexual characteristics. For convenience, small, intermediate, and large males are respectively referred to as Types 1, II, and III. A multi-year analysis of four study populations in east-central Florida revealed that males did not occur in discrete classes but, instead, occurred in a size and morphological continuum. Coloration and morphological development were highly correlated with size, and the mensural traits examined displayed strong allometry. Size distributions of mature males were relatively stable at any study site over time, but there were large differences between sites. At one extreme, males at the VABI site were always relatively small (16–40 mm SL) and Type III males were absent. At the other extreme, the population at the BDGST site contained a full array of male sizes (18–60 mm SL) and Type III males were present in small numbers throughout the summer months. Regression analysis revealed only minor differences between these extreme populations in the relationship between morphological development and body size. It appears that environmental, genetic, and social factors interact in a complex and poorly understood way to control variation in male size and morphology.     

Parallel evolution leads to reduced shoaling behavior in two cave dwelling populations of Atlantic mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, Poeciliidae,Teleostei)

Shoaling behavior protects fishes from avian and piscine predation, but at the same time costs of group living arise due to several mechanisms including increased food competition. Most cave fishes live in an environment in which avian and piscine predators are lacking, and cave environments are often characterized by low food availability, leading to increased food competition. Altogether, this should favor the reduction of shoaling in cave fishes. We compared shoaling behavior (i.e. the tendency to associate with a stimulus shoal) among surface dwelling populations of the Atlantic molly, Poecilia mexicana, and two cave forms of that species. The first cave population of P. mexicana originated from the Cueva del Azufre and was previously recognized as the only cave form of a poeciliid fish. The second cave population examined came from a cave that was discovered only recently (Cueva Luna Azufre). In both cave forms shoaling behavior was reduced compared with surface dwelling mollies.     

A new and morphologically distinct population of cavernicolous Poecilia mexicana (Poeciliidae: Teleostei)

The cave molly, Poecilia mexicana, from the Cueva del Azufre, a sulfur cave in Tabasco, Mexico, ranks among the best-studied cave fishes worldwide, despite being known from a single population only. Here we describe a newly discovered second population of cave-dwelling P. mexicana from a nearby, but mostly non-sulfidic cave (Luna Azufre). Despite apparent similarities between the two populations (such as reduced eye diameter and reduced pigmentation), a geometric morphometric analysis revealed pronounced morphological differentiation between the two cave forms.     

The role of female competition in the mating success of two clones of the Amazon molly, Poecilia formosa

Clones of the unisexual fish Poecilia formosa , the Amazon molly, live in the wild in association with their progenitor bisexual species, and compete with them for resources. These resources include food, space and also mates, since the Amazon molly reproduces gynogenetically, a process in which sperm transferred from a male host species activates their eggs. Earlier studies of the adaptive mechanisms that facilitate the coexistence of the bisexual and unisexual forms have focused upon male behaviour, but recent work with another unisexual species, Poeciliopsis , demonstrated that female aggressive behaviour may play an important part in determining the ecological standing of a clone.
We have extended our observations to female mating behaviour in two clones in the Amazon molly; clone 1 that does not thrive well under laboratory culture; and clone 2 that is hardy and survives very well. There was no aggression between the clones, nor did the clones show aggression towards females of the bisexual species. However, clone 1 consistently was more successful in the number of mating encounters made with the male; it approached the male more often than clone 2 and fled from the male less often. Males also showed some preference for mating with clone 1 females. If there are physiological differences in the wild clones similar to those seen in the laboratory, the mating advantage of the less robust clone 1 may ensure its continued survival in competition with a more vigorous clone.     

Interaction between female mating preferences and predation may explain the maintenance of rare males in the pentamorphic fish Poecilia parae

Variation in mating preferences coupled with selective predation may allow for the maintenance of alternative mating strategies. Males of the South American live‐bearing fish Poecilia parae fall in one of five discrete morphs: red, yellow, blue, stripe‐coloured tail (parae) and female mimic (immaculata). Field surveys indicate that the red and yellow morphs are the rarest and that their rarity is consistent across years. We explored the role of variable female mating preference and selective predation by visual predators in explaining the rarity of red and yellow males, and more generally, the maintenance of this extreme colour polymorphism. We presented wild‐caught P. parae females and Aequidens tetramerus, the most common cichlid predator, with the five male colour morphs in separate trials to determine mating and prey preferences, respectively. We found that a large proportion of females shared a strong preference for the rare carotenoid‐based red and yellow males, but a distinct group also preferred the blue and parae morphs. The cichlid predator strongly preferred red and yellow males as prey. Together, these results suggest that the interaction between premating sexual selection favouring and predation acting against the red and yellow morphs may explain their rarity in the wild. The trade‐off between sexual and natural selection, accompanied by variation in female mating preferences, may therefore facilitate the maintenance of the striking colour polymorphism in P. parae.     

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.     

Reproductive behavior of females and its relation to the mating success of two male forms inMnais pruinosa (Zygoptera: Calopterygidae)

The behavior of females of the damselflyMnais pruinosa was observed in the breeding season. Males consisted of both territorial (esakii) and non-territorial (strigata) types in the study area. Females sometimes arrived at a stream for mating and oviposition. The staying time of females in the stream was 1–3 hours, while that of males was 5–8 hours. The proportion of the males that copulated with the females did not differ between the two male forms present in the stream. In the Calopterygidae, of which the females usually perform multiple copulation, oviposition without subsequent recopulation is considered to be advantageous for a mated male in order to avoid the risk of sperm displacement. From such a viewpoint, the time spent for oviposition was measured for females that arrived at the stream after copulation with different male forms. The ratio was 69.1: 11.5–30.9 betweenesakii andstrigata males. Assuming that the oviposition time is proportional to the number of fertilized eggs laid, this ratio would represent the relative reproductive success of the two male forms. In fact, the relative abundance of the two male forms was also biased in favor ofesakii males (61.7:38.3) in this population. The mechanism of coexistence of the two male forms is discussed in relation to their reproductive success.     

Morphological and reproductive variation among populations of the Pacific molly Poecilia butleri

In viviparous organisms, pregnant females typically experience an increase in body mass and body volume. In this study, the prediction that variation in reproductive traits among populations of viviparous organisms should be related to variation among populations in body shape was tested in the Pacific molly Poecilia butleri, a viviparous fish that inhabits western Mexico and northern Central America. Variation among 10 populations in four reproductive traits was examined: brood size, individual embryo mass, total reproductive allotment and degree of maternal provisioning of nutrients to developing embryos. Variation among these populations in body shape was also examined. Significant variation among populations was observed in both brood size and reproductive allotment but not in embryo mass or degree of maternal provisioning. Significant variation among populations was also observed in body shape. After correcting for female size, however, reproductive traits and body shape were not associated among populations. This suggests that selective pressures acting on reproduction do not necessarily affect morphology and vice versa. Several factors might contribute to this unexpected lack of association between reproductive traits and morphology.     

圈养猞猁多雄交配雄性的成功繁殖

研究了笼养猞猁多雄交配的结果以及雄性的成功繁殖。雄兽与发情雌兽随机配对,其繁殖成功(所产生的后代数量)不取决于与雌兽的交配顺序、交配次数与持续时间、遗传关系或雄兽的行为特征,但似乎取决于精子质量,特别是取决于形态正常精子的百分率。繁殖成功的雄兽比其竞争对手具有更多的形态正常的精子,似乎更能成功地诱发交配雌兽的排卵。证实了4窝(占总窝数的20%)幼兽的双重父亲身份。在所有情形下, 2雄1雌的交配间隔为24h 。     

Male mating behavior and ejaculate expenditure under sperm competition risk in the eastern mosquitofish

Theory predicts that males should tailor the size of their ejaculatesaccording to temporal changes in the risk of sperm competition.Specifically, males are predicted to allocate more sperm toeach mating event with increasing risk (i.e., the probabilitythat the sperm from two males will compete for fertilization).We tested this hypothesis by using the eastern mosquitofish,a freshwater species of fish exhibiting a coercive mating systemand internal fertilization. We manipulated the perception ofsperm competition risk by adjusting the sex ratio under whichmales were maintained over 8 days. Males were housed eitherwith three females and one male (simulating high sperm competitionrisk) or with four females (low risk). After the treatment,we presented each test male individually to an unfamiliar male-deprivedfemale for 30 minutes and observed his mating behavior. We thenartificially stripped the test males of sperm and recoveredthe ejaculates from the females. Our results revealed that malesin the high-risk group performed higher levels of mating activityand sperm expenditure (i.e., used up more of their sperm reserves)than did low-risk males. A control experiment, in which testmales were treated but did not participate in the mating trials,revealed no significant difference in the number of sperm strippedfrom high- and low-risk males, indicating that sperm productionwas not affected by the treatment. We did not detect a differencein the number of sperm retrieved from females among the groups,raising the possibility that some sperm are lost during matingactivity, either through ejaculation with incomplete or interruptedpenetration, or via female ejection.     

Photophilic behaviour in surface- and cave-dwelling Atlantic mollies Poecilia mexicana (Poeciliidae)

The phototactic behaviour of surface- and cave-dwelling Atlantic mollies Poecilia mexicana was examined. Surface-dwelling Atlantic mollies, as well as light- and dark-reared cave fish showed photophilic behaviour under all light intensities used (620, 50 and 3 lx).     

The Effect of Gestational State on Oxygen Consumption and Response to Hypoxia in the Sailfin Molly, Poecilia latipinna

Metabolic activity in livebearing fishes increases with embryonic development so that embryos prior to parturition may have a higher mass-specific oxygen requirement than maternal tissues, temporarily increasing the total routine oxygen requirement of the female. We examined whether females of the livebearing poeciliid Poecilia latipinna (sailfin molly) increase their routine metabolic oxygen consumption during development of their broods. We also quantified effects of gestation on time allocation to aquatic surface respiration (ASR) under hypoxic conditions. Mass-adjusted routine metabolic rate (RMR) of female mollies showed a significant increase during late gestation. The RMR of males did not differ from females that were in their early or mid stage of gestation, but was lower than females in late gestation. Gestating females spent approximately 27% more time conducting ASR than non-gestating females when exposed to chronic hypoxia (1mgl^–1), further supporting a brood-related increase in oxygen demand. Increased time allocation to ASR may directly affect maternal predation risk in low-oxygen conditions.     

A Sexual Preference in the Amazon Molly, Poecilia formosa

The Amazon molly, Poecilia formosa, is an all-female species that reproduces by gynogenesis, i.e., it relies on sperm of males of closely related species to trigger embryogenesis. Sperm is supplied by males of P. latipinna and P. mexicana. Amazon mollies live in sympatry with at least one of these species, a few populations live in sympatry with two sperm-donor species. As P. formosa is sperm dependent, it needs mechanisms for species and mate recognition. To investigate the effect of rearing conditions on sexual preferences of Amazon mollies, we raised Amazon mollies in sympatry with P. latipinna and P. mexicana males. We used simultaneous choice tests to determine the effect of age on female sexual preferences. Immature Amazon mollies do not exhibit a preference if given a choice between a P. latipinna and a P. mexicana male, whereas adult P. formosa do have a preference for the P. latipinna male. We used two different stimuli in this study, live males and videotapes of males.     

Aspects of the mating behaviour of male mollies (Poecilia spp.)

The unisexual fish Poecilia formosa (the Amazon molly) reproduces by gynogenesis, a process in which sperm from the males of the host bisexual species activates development of its eggs. Unisexuals live with one of the host species in nature and compete with bisexual females for the males. It was long thought that male discrimination and mate selection established a balance between the unisexual and bisexual populations. Thus, hierarchies of males were set up in which dominant males mated with their conspecific females and subordinates mated with the Amazon molly. Recent evidence suggested, however, that male fish do not discriminate between their own females and the Amazon molly, and that there always are more males available than sexually receptive females of both species.
Our findings indicate that male behaviour may be more complex than suggested by either hypothesis. Mate discrimination and courtship behaviour appear to increase with age, so that large males show almost complete preference for their conspecific females, but smaller males will mate with the Amazon molly. In complex groups, small males often dart in and mate with their own females whilst the large males are engaged in courtship activities and defence of territory.