Ultraviolet-based female preferences in a viviparous fish

Studies on fish have demonstrated female-mating preferences based on dimorphic colours and colour patterns, but commonly focused on the 400-700 nm (visible) band. In experiment 1 we exposed females of the amarillo (Girardinichthys multiradiatus), a viviparous fish with visually based female mate choice, to two views of males either containing (UV+) or lacking (UV-) information in the 300-400 nm region. Experiments were conducted outdoors, and we found that females spent more time next to the male seen through the UV+ filter than the same seen through the UV- filter. Since the two images only differed in the content of UV radiation, we conclude that females of the amarillo can detect UV light. Experiments 2 and 3 were designed to test whether such an effect can be attributed to female preferences based on the males' UV-reflecting colour patterns. We first assessed which area(s) of the fish have UV-reflecting properties. Then, using an opaque gelatine powder to block skin reflectance, we found that, in the presence of environmental UV light, females spend more time close to the males with powder sprinkled outside the UV-reflecting area (control) than to the males with powder sprinkled over the UV-reflecting area (experiment 2). This could have been due to the concealment of signals in the visible spectrum, rather than in the UV spectrum; thus we repeated the same experiment indoors, in the absence of UV light (experiment 2b); here females failed to discriminate between males. Since we also found that females have UV-reflecting areas in their flanks, we conducted a third experiment to test whether the preferential association of females found in the first experiment was due to the tendency to school/associate with conspecifics, rather than to mate choice (experiment 3). We found no evidence of discrimination in this experiment. We conclude that the UV markings and vision in G. multiradiatus have a function in female mate choice. This is the first time that evidence has been found for the influence of UV for mate choice in fish.     

Inter-specific variation in anti-predator behavior in sympatric species of kangaroo rat

Kangaroo rats, Dipodomys, occupy desert habitats with little cover and thus are under high predation risk from diverse predators. The behavior used to assess predation risk or to escape capture is unknown. We therefore compared anti-predator behavior of two sympatric species of kangaroo rat of different sizes, D. merriami and D. spectabilis. We first examined whether kangaroo rats use olfaction as a first line of defense against snake predation and tested the rats for their responses to scent extracted from two species of snake that live sympatrically with the kangaroo rats, the Mojave rattlesnake (Crotalus scutulatus) and the gopher snake (Pitophis melanoleucus). We also tested for species differences in anti-predator behavior through 15-min interactions between the kangaroo rats and free-moving gopher snakes. We found that D. spectabilis actively approached the scent of both rattlesnakes and gopher snakes more than controls of vegetable oil and evaporated solvent (Fig. 1). In contrast, D. merriami did not differentiate snake odors from controls in the experimental arena, but they sniffed the sand where a free-moving snake had passed more than D. spectabilis. Both species successfully avoided predation in encounters with live snakes. Although total numbers of approaches and withdrawals were similar (Fig. 2), D. spectabilis spent significantly more time within striking distance of the snake than D. merriami. D. spectabilis approached the head of the snake in 93% of its approaches and often engaged in nose to snout contact with the snake. If the snake struck, D. spectabilis jumped directly backward to avoid a strike and footdrummed at a safe distance. In contrast, D. merriami oriented to the snake more than D. spectabilis, but approached the head in only 41% of the approaches and rarely engaged in nose-to-snout contact. The snakes struck, hissed and decreased predatory approaches with D. spectabilis but not with D. merriami (Fig. 3). These results show that kangaroo rats can behaviorally influence the risk of being preyed on by snakes. The two species differ, however, in how they react to snakes. The larger D. spectabilis confronts snakes while the smaller D. merriami monitors snakes from a safe distance and avoids them.     

Least auklet ornaments: do they function as quality indicators?

Summary In the monogamous least auklet (Aethia pusilla, Alcidae) both males and females have three highly variable ornamental traits (facial plumes, a colourful bill and a knob-like bill ornament) and both sexes perform courtship displays. To assess whether mating preferences could be related to the expression of these ornaments, we performed model presentation experiments in which we varied the bill colour and the size of both the bill ornament and the facial plumes. Auklets reacted to models that had brighter red bills and accentuated facial plumes with more frequent sexual displays than to models with average bills and plumes. We conclude that these two ornamental traits are likely to be favoured by sexual selection through mating preferences. In general, however, ornaments were weak predictors of individual quality. Multiple regression indicated that all ornaments taken together explained a small (R ² = 0.07) but significant proportion of the variability in adult body condition but only in a poor year for reproduction when the birds were in relatively poor body condition and a small proportion of the population bred successfully. The degree of ornamentation was also not related to timing of breeding, chick feeding rate or reproductive success and there was no relationship between adult survival and ornaments. We conclude, therefore, that least auklet ornaments are, at best, weak indicators of quality that may be the result of sexual selection operating only in years when breeding conditions are poor. Offprint requests to: R. Montgomerie     

Courting male garter snakes (Thamnophis sirtalis parietalis) use multiple cues to identify potential mates

Mating aggregations of red-sided garter snakes (Thamnophis sirtalis parietalis) in Manitoba provide a unique opportunity to identify the cues that attract a male snake and induce him to court. The snakes are abundant, tolerate human presence, and males direct courtship to a subset of other males ("she-males") as well as to females. Previous work has emphasised the role of pheromonal cues (skin lipid profiles) as sexual attractants in snakes. However, pheromones are so widely distributed throughout the den area that these chemical cues may be difficult for males to use to localise females. Our field studies show that males and females differ in several other attributes such as size (females are larger than males), body temperature (very cold snakes are often females), muddiness (females are often covered in mud) and whether or not they are solitary (most females are surrounded by courting males). Experiments show that males use all of these cues to identify possible sexual partners. Visual and thermal cues are particularly important for solitary mate-searching males; but after a mating ball forms around a female, pheromones may be most important. Our study also reveals substantial variation among individuals: females vary in attractiveness, and males vary in their intensity of courtship as well as in the magnitude of their preference for one potential partner versus another. Thus, a male snake's "decision" whom to court depends not only on visual and thermal as well as chemical cues, but also on the male's own preferences and on subtle differences among potential sexual "targets".     

A cost worth paying: energetically expensive interactions with males protect females from intrasexual aggression

When approached by males, females of the Amarillo fish (Girardinichthys multiradiatus) perform a behaviour called vibration or they are aggressively challenged. We quantified vibration and assessed whether it compromises the rate of feeding attempts in dyads kept in outdoor enclosures. Male approaches resulted in female vibration and in a reduced feeding rate. Vibration was not evoked by female–female aggression, which was frequent and always ended in the subordinate fleeing from the dominant female. Using a closed respirometer we found that vibration is costly; oxygen consumption of females was greater in the presence of a male (which evoked vibration) than in the presence of a non-familiar female (when no vibration occurred). By recording interactions of females confined in aquaria in the presence and in the absence of males, we confirmed that escaping is the only available response to deal with female aggression. Females kept without males participated in frequent aggressive (even lethal) interactions that did not abate while the subordinate female was in sight of the dominant, and which caused premature births and injuries. Yet in the alternative treatment aggression ceased when a male approached, prompting vibration in both females. Thus, in the Amarillo, in as much as it evokes energetically costly female vibrations, male courtship is an expression of sexual conflict. However, in the absence of males, frequent female aggression potentially annuls the benefits of not vibrating. We propose that a complete appraisal of the consequences of sexual conflict must include an assessment of the costs imposed by intra-sexual interactions.     

Congenital predispositions and early social experience determine the courtship patterns of males of the Amarillo fish

Mounting evidence indicates that learning shapes fish foraging and social behavior, which suggests that it may also underlie the frequently reported geographic variation in courtship. In this study, we analyzed how early social experience influences courtship patterns in males of the Amarillo fish (Girardinichthys multiradiatus). In a first experiment, we quantified the courtship of males from two allopatric populations grown in isolation and estimated the heritability of a key attribute of their courtship patterns. Then, we raised males from two allopatric populations with either sympatric or allopatric females and at two different densities, and registered their adult courtship patterns towards females from a third population. We found that the courtship pattern of males has a significant heritable component, yet in the second experiment, it was influenced by what type of females males were raised with, by their social rank (dominant/subordinate/singleton), and by the behavior of the female that they were courting. We conclude that courtship behavior in this fish has a congenital predisposition and is affected during development by different aspects of early experience and seemingly remains flexible throughout life; a complex ontogenetic trajectory that under natural conditions may have led to the locally stable dialect types observed in the courtship patterns of males from allopatric populations.     

Female mallard mating preferences for multiple male ornaments

Mating preferences were studied in captive wild-stock female mallards. Independent observers scored ornament brightness for the bill and eight plumage areas of male mallards on a scale of 1 to 6. Ornament scores were generally positively correlated with each other as indicated by univariate correlations and principal component analysis. Males with higher bill scores were significantly more likely to obtain pairings. None of eight individual plumage ornaments were significant predictors of pairing success, but males with higher average plumage scores had significantly higher pairing success. Males that completed the molt first were also significantly more likely to be paired. Thus females may have paid some attention to plumage brightness. Stepwise multiple regression produced a model that included only bill and molt, both of which contributed a similar degree (r ² = 0.39); plumage ornaments and size measurements did not enter the model. These results are interpreted in light of hypotheses for the evolution of multiple ornaments, and phylogenetic plumage patterns in dabbling ducks. Received: 18 December 1995 / Accepted after revision: 22 September 1996     

Effects of reproduction on the antipredator tactics of snakes (Tropidonophis mairii, Colubridae)

Does an animal’s reproductive state influence the distance at which it flees from an approaching predator? We predicted that reproduction would increase approach distances in pregnant females (because they are burdened with eggs and thus less able to escape rapidly), but reduce them in males (because of lower vigilance due to males focusing on mate-searching rather than predator detection). Field data on approach distances of keelback snakes (Tropidonophis mairii, Natricinae, Colubridae) supported both of these predictions. We walked the same 1.3-km transect along a dam wall in tropical Australia on 135 nights, and recorded the distances at which snakes fled from our approach. Locomotor speeds were measured for a subset of these animals. Variations in approach distance due to season, weather conditions, prior capture history and snake body size were minor, but reproduction strongly affected approach distances for snakes of both sexes. Gravid females were slower than non-gravid females, and fled at greater distances. Reproductive status did not affect locomotor speeds of males, but males that were reproductive (i.e., contained sperm) permitted closer approach than did non-reproductive adult males. Reproduction thus affected approach distances in snakes of both sexes, but in opposite directions and for different reasons.     

Chemical ornaments of male lizards <Emphasis Type="Italic">Psammodromus algirus</Emphasis> may reveal their parasite load and health state to females

Evidence for parasite-mediated sexual selection has been found in many species that use visual ornaments to attract females. However, in many animals, variation in female responses to scents of parasitized males suggests that parasitic infections might also affect information conveyed by pheromones (i.e., chemical ornaments). Thus, pheromones might also function in parasite-mediated sexual selection. We show here that female lizards Psammodromus algirus responded differently to femoral gland secretions of males according to the parasite load and health of these males. Scents of healthier males elicited more tongue flicks (a chemosensory behavior) by females, suggesting that these scents were more attractive. Chemical analyses showed that parasite load and the T-cell-mediated immune response were related to the variability in the proportions of some lipids in secretions of males. Further trials testing the chemosensory responses of females to chemical standards indicated that females actually discriminated the chemicals related to males’ health from other chemicals found in secretions. We suggest that these chemical ornaments may provide reliable information on the health and degree of parasitic infection of a male.     

Female preferences for male calling bout duration in a field cricket

Summary The calls of male field crickets attract sexually receptive females. In Gryllus integer, males differ from one another in their durations of uninterrupted calling (calling bout lengths). Tape recordings of the calls of 50 wild-caught males revealed that 14 males spent most of their calling time in short bouts (Fig. 1A), 18 in both short and long bouts (Fig. 1B), and 18 in long bouts (Fig. 1C). Re-recordings of 32 males after 3 weeks showed that calling bout lengths of individual males are stable with time (age) (Fig. 2). Three phonotaxis experiments investigated whether calling bout lengths of males affect female preferences. They demonstrated that (1) females can discriminate among conspecific males on the basis of calls alone; (2) females are preferentially attracted to males with long calling bout lengths; and (3) calling bout length is the specific factor responsible for preferential attraction. These results precisely identify a criterion that females use to discriminate among potential mates of their own species.     

Sexual conflict in the snake den

Red-sided garter snakes (Thamnophis sirtalis parietalis) court and mate in spring, soon after they emerge from large communal overwintering dens in south-central Manitoba. Because of a massive bias in the operational sex ratio, every female attracts intense courtship from dozens to hundreds of males. We suggest that this courtship constitutes significant ”harassment,” because it delays the females’ dispersal from the den and hence increases their vulnerability to predation. Small females may face the greatest costs, because they are less able to escape from amorous males (who court all females, even juvenile animals). Our measurements show that males are stronger and faster than females. Experimental trials confirm that the locomotor ability of females (especially small females) is greatly reduced by the weight of a courting male. Arena trials show that intense courtship stimulates females to attempt to escape. Remarkably, some females that are too small to produce offspring may nonetheless copulate. This precocious sexual receptivity may benefit juvenile females because copulation renders them unattractive to males, and thus allows them to escape more easily from the den. Female ”tactics” to escape male harassment may explain other puzzling aspects of garter snake biology including size-assortative mating, temporal patterns in dispersal from the den, avoidance of communal dens by young-of-the-year snakes, and female mimicry. Hence, sexual conflict may have influenced important features of the mating system and behavioral ecology of these animals. Received: 8 May 2000 / Revised: 28 July 2000 / Accepted: 30 July 2000     

Does the cost of polygyny in house wrens include reduced male assistance in defending offspring?

Summary This study investigated whether reduced male aid in defending offspring potentially reduces the fitness of females choosing already-mated males in the house wren (Troglodytes aedon), a small, territorial songbird. Frozen snakes were placed at 23 nests of monogamously mated males and 12 secondary nests of bigamously mated males. All presentations were made during incubation stages of females attending focal nests. Snakes were placed at nests of secondary females when nests of their primary counterparts contained young 5–9 days old. Males are most attentive to primary nests during this period and should therefore be relatively inattentive to secondary mates and nests. Nevertheless, an equal proportion of monogamous and bigamous males discovered snakes within 15 min, and mean time to discovery, when discovery occurred, did not differ with nest status. Monogamous and bigamous males were also equally likely to attack snakes physically once discovered. Monogamous males appeared no more likely to discover snakes than bigamous males for two main reasons. First, although monogamous males were near focal nests (i.e., < 10 m) more often than bigamous males, monogamous males tended to stay out of view of nests for long periods. In contrast, bigamous males always went immediately to focal nests upon arriving in their vicinity. Second, about one-third of monogamous males in this study spent much of their time during trials at the far edges of their territories advertising for secondary mates. Our experiment suggests that reduced male aid in defending nests against small, diurnal predators probably does not contribute to the cost of polygyny in house wrens. Correspondence to: L.S. Johnson     

Male mate choice favors more colorful females in the gift-giving cabbage butterfly

While the phenomenon of male mate choice has attracted considerable attention in the last two decades, whether this sexual selection mechanism could drive the evolution of female ornaments remains poorly understood. Here, we used experimental manipulation of female wing coloration to investigate male mate choice in Pieris rapae, a gift-giving butterfly. Further, we tested whether males’ nutritional status influenced their mating preferences by subjecting larvae to short periods of starvation. We found that males showed significantly more mating approaches toward control females with more colorful wings (higher pteridine content), and that this preference was strongest in low-nutrition males. Additionally, a study of field-collected females revealed that pteridine-based wing coloration was positively correlated with female egg load, which suggests such ornaments may signal female quality. Pteridine-based ornaments are widespread in nature, however their potential as honest signals in male mate choice remains largely unexplored. This work furthers our understanding of how male mate choice and female ornamentation may evolve in species whose mating systems include nutritional nuptial gifts.     

Sea Snake Harvest in the Gulf of Thailand

Conservation of sea snakes is virtually nonexistent in Asia, and its role in human–snake interactions in terms of catch, trade, and snakebites as an occupational hazard is mostly unexplored. We collected data on sea snake landings from the Gulf of Thailand, a hotspot for sea snake harvest by squid fishers operating out of the ports of Song Doc and Khanh Hoi, Ca Mau Province, Vietnam. The data were collected during documentation of the steps of the trading process and through interviewers with participants in the trade. Squid vessels return to ports once per lunar synodic cycle and fishers sell snakes to merchants who sort, package, and ship the snakes to various destinations in Vietnam and China for human consumption and as a source of traditional remedies. Annually, 82 t, roughly equal to 225,500 individuals, of live sea snakes are brought to ports. To our knowledge, this rate of harvest constitutes one of the largest venomous snake and marine reptile harvest activities in the world today. Lapemis curtus and Hydrophis cyanocinctus constituted about 85% of the snake biomass, and Acalyptophis peronii, Aipysurus eydouxii, Hydrophis atriceps, H. belcheri, H. lamberti, and H. ornatus made up the remainder. Our results establish a quantitative baseline for characteristics of catch, trade, and uses of sea snakes. Other key observations include the timing of the trade to the lunar cycle, a decline of sea snakes harvested over the study period (approximately 30% decline in mass over 4 years), and the treatment of sea snake bites with rhinoceros horn. Emerging markets in Southeast Asia drive the harvest of venomous sea snakes in the Gulf of Thailand and sea snake bites present a potentially lethal occupational hazard. We call for implementation of monitoring programs to further address the conservation implications of this large‐scale marine reptile exploitation. Cosecha de Serpientes Marinas en el Golfo de Tailandia     

Does promiscuous mating preclude female choice? Female sexual strategies in chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d’Ivoire

According to sexual selection theory, females should selectively mate with high-quality males to enhance offspring survival and maximize reproductive success. Yet, chimpanzee (Pan troglodytes) females are known to mate promiscuously. Although there is substantial rationale for a promiscuous mating strategy, there is also a strong expectation that females should be selective, and the question arises as to whether promiscuity precludes female choice. The aims of this study are to: (1) compare wild female chimpanzee sexual strategies throughout estrus, and (2) determine whether females exhibit mate preferences for particular males. Over 2,600 h of data were collected on two habituated chimpanzee (Pan troglodytes verus) communities in the Taï National Park, Côte dIvoire. Female mate preferences were measured by quantifying proceptivity and receptivity toward males. Results indicate that all females exhibited proceptivity and resistance to male solicitations, but that there was substantial variation in their magnitudes within and among females. Female proceptivity rates were lower and resistance rates were higher in the periovulatory period (POP) when conception is most likely. Females were more selective during POP, and more promiscuous outside of POP, suggesting that females may follow a mixed reproductive strategy, being selective when conception is likely and more promiscuous when conception is unlikely. Results from this study emphasize the importance of considering the fertility window when determining female mate preferences, and of examining female behavior in POP and non-POP phases separately when evaluating hypotheses for multi-male mating.Communicated by C. Nunn     

Population differences in how black-tailed prairie dogs deal with snakes

Summary This paper examines the antisnake behavior of a snake-experienced Texas (TX) population and a snake-naive South Dakota (SD) population of black-tailed prairie dogs (Cynomys ludovicianus). Animals in both populations were presented with tethered rattlesnakes and nonvenomous bullsnakes before and after the first emergence of pups from their natal burrows. SD and TX adults were more likely to actively harass snakes than were pups. There were no differences in the behavior of SD adult males vs females or fathers vs non-fathers, contrary to what was found in TX. However, SD mothers did call more and stayed closer to snakes than did SD non-mothers. All SD adult sex-parental classes spent less time dealing with snakes after the emergence of pups, which was not observed in TX. The primary difference between the two populations was the behavior of TX males who spent a great deal of time harassing snakes. SD adults behaved most like TX females in that these groups spent little time actively dealing with snakes. Pups in both populations behaved similarly. These results are interpreted in terms of the relationship between potential predator and potential prey.     

Space use and social structure in meadow voles,Microtus pennsylvanicus

Summary Free-ranging, sexually mature meadow voles (Microtus pennsylvanicus) were tracked by using radiotelemetry from June through August in Front Royal, Virginia, U.S.A. Estimates of intraspecific spacing were derived from the concurrent movements of up to 16 voles. Positions were recorded hourly for 24 h, twice per week. A total of 16 male and 15 female voles were studied during sixteen 24-h sessions.The daily ranges of males (192.3±109.7 m²) were larger and more variable than those of females (68.6±39.4 m²). Males also changed locations more frequently (Fig. 2).Adult females usually maintained territories free of other females; males overlapped considerably among themselves (Fig. 2). Males temporarily moved into the areas occupied by estrous females, indicating intrasexual competition among males for access to receptive females (Fig. 3). M. pennsylvanicus appears to be promiscuous, is socially organized into territorial, maternal-young units during the breeding season, and fits the female territorial model of population regulation.     

Female choice of a non-bodily ornament: an experimental study of cichlid sand craters in Cyathopharynx furcifer

In many species of East African cichlids, males build sand craters or “bowers” to attract females for spawning. It is commonly assumed that these exaggerated sexual traits have a function similar to bodily ornaments. In non-bodily ornaments, however, the behaviour creating the structure may serve as an additional source of information to potential partners, providing multiple signals for mate choice. We tested whether and to what extent females use these signals for choosing males by observing the individual sampling behaviour of female Cyathopharynx furcifer in the field. In addition, we experimentally manipulated crater characteristics in the field and laboratory. We found that females spawn preferably with owners of large and well-maintained craters, but when crater size was enlarged or reduced, the resulting building activity of crater owners affected female choice more strongly than the manipulated crater size per se. We discuss the importance of multiple signals in species constructing extended phenotypes.     

Facultative pheromonal mimicry in snakes: “she-males” attract courtship only when it is useful

Males of many animal species mimic females, and thereby deceive rival males. Facultative shifts in posture, color, or movement allow a male using visually-based mimicry to adopt and terminate mimicry rapidly. Pheromonal mimicry is rare in vertebrates perhaps because it is difficult to redeploy pheromones rapidly enough to adjust male tactics to local conditions. In Manitoba garter snakes (Thamnophis sirtalis parietalis), female mimicry benefits males immediately after they have emerged from hibernation. While the snakes are cold and slow, courtship warms them and protects them against predatory crows. This benefit disappears as soon as the snakes are warm. We show that (unlike females) she-male garter snakes attract courting males only when they are cold. Low temperatures may suppress volatility of “less attractive” components of the pheromones (saturated methyl ketones) that she-males use to attract courtship, allowing male snakes to function as transvestites only when this tactic is beneficial.     

Do female association preferences predict the likelihood of reproduction?

Sexual selection acting on male traits through female mate choice is commonly inferred from female association preferences in dichotomous mate choice experiments. However, there are surprisingly few empirical demonstrations that such association preferences predict the likelihood of females reproducing with a particular male. This information is essential to confirm association preferences as good predictors of mate choice. We used green swordtails (Xiphophorus helleri) to test whether association preferences predict the likelihood of a female reproducing with a male. Females were tested for a preference for long- or short-sworded males in a standard dichotomous choice experiment and then allowed free access to either their preferred or non-preferred male. If females subsequently failed to produce fry, they were provided a second unfamiliar male with similar sword length to the first male. Females were more likely to reproduce with preferred than non-preferred males, but for those that reproduced, neither the status (preferred/non-preferred) nor the sword length (long/short) of the male had an effect on brood size or relative investment in growth by the female. There was no overall preference based on sword length in this study, but male sword length did affect likelihood of reproduction, with females more likely to reproduce with long- than short-sworded males (independent of preference for such males in earlier choice tests). These results suggest that female association preferences are good indicators of female mate choice but that ornament characteristics of the male are also important.