The reproductive behavior of the longspine snipefish,Macrorhamphosus scolopax (Syngnathiformes,Macrorhamphosidae)

Synopsis About two hundred Macrorhamphosus scolopax were observed in a large tank and their behavior was videotaped for subsequent analysis. When not disturbed, the fish moved slowly in a head-down position. Spawning in the tank took place in the last two hours before dark. Courting males change color and actively interact aggressively with other males. They push against each other while swimming in parallel or they charge their opponents with their erected dorsal spine directed at them. Courtship begins near the bottom. After following and swimming parallel to the female the two fish unite tightly by their caudal peduncle, with their ventral regions in close proximity and rise slowly in the water column. During the ascent, the genital papilla of the female extends repeatedly and contacts the genital region of the male.     

The social and reproductive behavior of the eyed flounder,Bothus ocellatus,with notes on the spawning ofBothus lunatus andBothus ellipticus

Synopsis Observations of the Caribbean eyed flounder,Bothus ocellatus, were made during two field trips to Bonaire, Netherlands Antilles (January 1990, and December 1990 – January 1991) at depths from 3 to 6 meters. Data were collected by six scuba divers during 54 dives totaling 70 hours. Three mating groups were observed, each consisted of a male and 1 to 6 females. Each female had a distinct subunit within the male's territory. Courtship activity began about one hour before sunset and continued until sunset, when the male retired for the night. Both males and females retired into the sand at locations outside their daytime territory. Every day, males attempted to mate with each female individually within their territory. Spawning began with the male moving under the female who was resting on the sandy bottom. The pair then made a slow upward rise approximately 15 to 75 cm above the substrate, which culminated in a release of a cloud of gametes.Bothus lunatus andBothus ellipticus exhibited similar social organization toB. ocellatus, but differences in the spawning rise were observed. Intraspecific agonistic behavior was observed within males ofB. ocellatus andB. ellipticus.Senior author     

Androgen and the elaborate courtship behavior of a tropical lekking bird

In most bird species, male courtship behavior is controlled by testosterone (T) and its metabolites. In species breeding in temperate and arctic regions T circulates at high levels during a relatively short courtship period because high levels of T can be costly in terms of immunocompetence and parental care. Few studies have investigated androgen modulation of courtship behavior in tropical birds. Male golden-collared manakins (Manacus vitellinus) aggregate in leks for several months and perform spectacular, acrobatic courtship displays. Here we examined whether T is elevated in golden-collared manakins during the displaying period and if courtship behavior is modulated by androgen action on androgen receptors. We measured T levels in displaying males at the beginning of the breeding season and again, one month later. In addition, both wild and captive males were treated with the anti-androgen, flutamide, and their courtship behavior was recorded for several weeks. T levels were relatively high shortly after leks were established but decreased substantially a month later, even though the amount of courtship did not change. Flutamide reduced male courtship activity for one week, but display behavior then increased after two weeks of flutamide treatment. Our studies show that androgens modulate male manakin courtship, but the amount of courtship is not directly correlated with the concentration of circulating T. These results suggest that the relationships between androgen and courtship might differ between tropical and temperate birds.     

Agonism,ranking, and the social behavior of adult male baboons

An ingrained technique in primatological analysis is to rank individual animals in a hierarchical system that is usually but not always based on the outcome of agonistic encounters. An explication of group structre and function is then sought by correlation these rankings with aspects of behavior. When the behavior of the adult males in a free-ranging troop of olive baboons is examined, ranking is found to be inadequate to represent the behavioral variability existing between individuals. The relationships maintained by these animals is analyzed, emphasizing amounts of time devoted to specific activities, and it is recommended that affiliative rather than agonistic behavior be stressed in future studies.     

Courtship behavior of male white perch, Morone americana: evidence for control by androgens

Courtship behaviors are androgen-dependent in many vertebrates and castration often decreases courtship. We examined the effectiveness of castration in reducing courtship behaviors and 11-ketotestosterone (KT) and testosterone (T) in restoring them in male white perch. Castrates were given implants containing KT, T or no hormone. Sham-operated males received implants without hormone. Three weeks later, males were exposed to an ovulated female for 1 h and two courtship behaviors were quantified. Attending behavior involves close and continuous following of a female with occasional contact. Circling involves rapid transits around the female in a circular pattern or back and forth in front of her. In plasma samples taken immediately after observations, KT and T were below detectable levels in castrated males but at high physiological levels in males implanted with KT or T. Castrated males given KT attended females more than castrated males given T implants or implants containing no hormone, but not more than sham-operated males. Circling was eliminated by castration but restored by implantation with T or 11-KT to values exhibited by sham-operated males. This is one of the few demonstrations that KT can regulate courtship behavior in a non-territorial and economically important fish species.     

Reproductive behavior of the sergeant major,Abudefduf saxatilis,within a closed system aquarium

Synopsis The reproductive and social biology of the sergeant major,Abudefduf saxatilis, was studied for six months in The Living Seas aquarium (the largest aquarium ever constructed). The purpose of the study was to discern possible behavioral differences between captive and wild breeding populations of this species. A total of 350 h of observation were obtained using closed circuit SCUBA, open circuit SCUBA, and remote operated vehicles. Slight differences were observed in the behavior of individuals in the sample population and those seen in wild populations. These differences may be due to the effects of a controlled closed system on the biology of these fishes.     

Reproductive behavior and social organization in the sand tilefish,Malacanthus plumieri

Synopsis Observations on the Caribbean sand tilefish, Malacanthus plumieri, were made during 3 dive trips in the Bahamas (Feb., 1986; July, 1986; Jan., 1987) at depths from 5 to 30 m. Eighteen mating groups contained 1 to 5 females (x = 2.9). Seven male home ranges (encompassing female ranges within mating groups) measured 250 to 700 m². Of 134 paired rises recorded between 1733 and 1925 h EST, 38 had an observed cloud of gametes. Mating times are presented in relation to time of sunset and depth of water. Courtship begins with conspicuous looping behavior of the male as he swims to a female. Close pairing, criss-crossing, and low rises (<1.5 m) often precede a high quivering upward swim (1.5 to 6 m) which culminates in the release of a cloud of gametes at the apex. M. plumieri is compared to the protogynous hermaphroditic razorfishes, Xyrichtys pentadactylus and X. martinicensis, with regard to social organization, size of male home ranges in exposed sandy habitats near coral reefs, sexual size dimorphism, and sex ratio.     

Sexual selection,reproductive behavior,and speciation in the mbuna species flock of Lake Malawi (Pisces: Cichlidae)

Synopsis The cichlid species flocks of the African Great Lakes represent the most extreme case of adaptive radiation among vertebrates. Recently, attention has focused on the potential for sexual selection to drive or accelerate speciation in these fishes. Cichlids as a whole are social in nature and display complex behavior, particularly during courtship and spawning; however, the extent to which changes in species recognition cues may account for species diversity among haplochromine lineages has remained speculative. Our investigations have indicated that oral incubating haplochromines show a reduction in diversity and extent of courtship relative to substrate brooding cichlids, and apparently retain aspects of a primitive specific mate recognition system. Laboratory observations of courtship in the Malawian endemicPseudotropheus zebra suggest that organization of the spawning bout is loose, and lacking in any well defined stimulus-response chain. Interspecific comparisons of in situ courtship behavior among male mbuna, lithophilous haplochromines of Lake Malawi, revealed only one potential example of species-specific behavior, and indicated that mate choice occurred prior to the onset of intense courtship. Courtship display by male mbuna does not appear critical to species recognition and may represent an evolutionary relict. Alternatively, male courtship display may have an indirect effect on successful reproduction or may be maintained through pleiotropic effects. The mbuna display no evidence of behavioral innovation and show limited interspecific differentiation in behavioral expression. More generally, there is no evidence at present to suggest that epigamic sexual selection, acting on courtship behavior, has been a major mechanism in the diversification of the haplochromine species flocks.     

Female sexual swelling size, timing of ovulation, and male behavior in wild West African chimpanzees

Conspicuous swellings of the perineal skin can be observed in females of many catharrine primate species particularly during the middle stages of the ovarian cycle. The functional significance of this trait remains poorly understood. Recently, two hypotheses, the "reliable indicator" hypothesis and the "graded signal" hypothesis that take into account not only the pattern but also the exaggerated size of sexual swellings, have gained attention. Here we test several predictions made by these hypotheses by combining (i) direct size measures (from video captures) of female sexual swellings with (ii) urinary hormone data to indicate timing of ovulation through enzyme immunoassay measurements of estrone conjugates and pregnanediol glucuronide (PdG) and (iii) behavioral observations of male mating efforts throughout 36 ovulatory cycles in 12 wild chimpanzees. We are able to show that (i) even within the traditionally defined maximum swelling period, further slight increases in swelling size indicate approaching ovulation, and (ii) that male mating interest changes according to the changes in swelling size. Furthermore, absolute swelling size during the periovulatory period increases and the alpha male associates more with females as the number of cycles to conception decreases. Finally, when having the choice between several "maximally" tumescent females, the alpha male prefers the female that is in the fertile phase of her cycle rather than that with the biggest swelling at that time. Thus, most of our findings are in line with the predictions of the graded signal hypothesis while none of them would support the reliable indicator hypothesis.     

Stereotyped sequence of mating behavior in the Far Eastern catfish, Silurus asotus, from Lake Biwa

 Mating behavior of the Far Eastern catfish, Silurus asotus (Siluriformes: Siluridae), was observed in a ricefield system facing the shore of Lake Biwa in mid-May to early June in 1990–1997. A set behavioral sequence similar to those of two other silurid fishes, S. biwaensis and S. lithophilus, both endemic to the Lake Biwa system, was observed: “chasing,”“clinging,”“enfolding” while “squeezing” by the male, and “circling” by the spawning pair. This form of mating behavior is quite different from that of S. asotus reported from the Ooi River system in Kyoto Prefecture, which mainly spawns in running water in ditches. Received: April 10, 2001 / Revised: November 5, 2001 / Accetped: November 20, 2001     

Non-verbal behavior as courtship signals: the role of control and choice in selecting partners

In this work, we provide evidence based on direct observation of behavior in encounters of opposite-sexed strangers, that women initiate and “control” the outcome. In the first minute of these videotaped 10-min interactions, neither female “solicitation” behavior nor “negative” behavior is strongly related to professed interest in the man, while female “affirmative” behavior at this stage modulates male verbal output in later stages (4–10 min). Although the rate of female courtship-like behavior is significantly higher in the first minute, it is only in the fourth to tenth minute that the rate of female courtship-like behavior is correlated with professed female interest. We hypothesize that this serves as a strategic dynamic reflecting sexual asymmetry in parental investment and the potential cost of male deception to women. Ambiguous protean behavioral strategies veil individuals' intentions and make their future actions unpredictable. These behavioral strategies may result in men's overestimation of female sexual interest.     

Testosterone, reproductive stage, and territorial behavior of male and female European stonechats Saxicola torquata

We investigated territorial behavior and circulating testosterone (T) levels in a multiple-brooded population of the European stonechat, a socially monogamous passerine bird with biparental care. Between arrival at and departure from the breeding territories, we (1) quantified behavior of both sexes in response to a simulated territorial intrusion (STI) of a male conspecific and (2) measured plasma T concentrations in males and females. Male response scores to a STI and male T concentrations varied with phase, but there was no temporal association between plasma T levels and the intensity of territorial behavior. During both two sexual and two parental phases, at least half of the tested males showed aggressive responses. About 20% of the tested males responded with courtship prior to laying of the first clutch, but none of the males courted during later phases. Age had a positive overall effect on male plasma T. Females also reacted to the STI of a male, but their responses did not vary with breeding phase. Female plasma T varied with phase, being elevated during production of the first but not of the second clutch. As with males, female responses to the STI were not correlated with T levels. Responses of pair partners were positively correlated with each other. We conclude that modulation of male territorial aggression with breeding phase is not regulated by changes of plasma T concentrations. In light of other studies showing reduced male aggression by pharmacological inhibition of cellular actions of T, we propose that T is permissive for male territorial aggression, but does not mediate short-term changes associated with breeding phase. The function of the high female plasma T concentrations during formation of first clutches could be related to the production of eggs with high concentrations of androgens.     

Sociosexual behavior, male mating tactics, and the reproductive cycle of giraffe Giraffa camelopardalis

Female distribution exerts a major impact on male mating tactics. Giraffe cows have a reproductive cycle, and a social system, that should favor a male roaming reproductive tactic. We conducted a 2-year study of female Rothschild's giraffe (G. c. rothschildi) reproductive endocrinology in order to characterize attributes of the reproductive cycle and investigate how female endocrine and behavioral cues influence mating activity. We used non-invasive fecal steroid methods to determine reproductive state among females residing in a herd in a large outdoor enclosure. We found that females had an estrous cycle of 14.7 days and that they regularly had multiple ovarian cycles prior to conception. Adult males were more likely to associate with, and sexually investigate, females when they were cycling than when they were either pregnant or acyclic. During the estrous cycle, male-female proximity and sociosexual behavior were more pronounced during the probable fertile phase than the rest of the cycle. Sexual activity between giraffe coincided with the periovulatory period, with male interest in females peaking during the fertile window in the absence of proceptive behavior by females. We conclude that males detect reliable cues revealing female reproductive status and partition their reproductive effort in response to such cues. We propose that male giraffe adopt a roaming reproductive strategy with their large size, enabling them to search for and mate guard fertile females while minimizing metabolic costs.     

Spawning and nesting behavior of the waccamaw darter,Etheostoma perlongum

Synopsis The spawning and nesting behavior ofEtheostoma (Boleosoma) perlongum was investigated in the field and laboratory. Sexual dimorphism is highly developed in such features as genital papillae, first dorsal and paired fins, and nuptial coloration. A reproductive migration from mid-lake to shore occurs in the spring: males precede females to select nest sites under submerged sticks and other debris. The male excavates a depression beneath the submerged object. Gonad analysis indicates a single spawning season extending from March through June. Nests were found from late April to mid-June and were guarded by a single male for periods of 13 to 36 days. Males initiate courtship by lateral display, lead the female to the nest site and show the nest by inverting. The female responds by tail up, tail wag and circle; males also tail wag and circle. Spawning pairs invert, usually in unison, and orient head to head or, less often, head to tail. The female deposits eggs while holding her body in a weak S or J shape with the caudal peduncle held away from the spawning substrate while vibrating.     

Seasonal changes in territoriality and frequency of agonistic behavior in two densities of juvenile brown bullhead,Ictalurus nebulosus

Synopsis Incidence of territoriality and frequency of six agonistic action patterns of juvenile brown bullhead, Ictalurus nebulosus, were observed in the laboratory over a two year period at two fish densities under conditions of constant light and temperature. High density fish showed little territoriality or agonistic behavior while low density fish displayed an annual cycle of agonistic behavior and territoriality. Agonistic behavior frequency and incidence of territoriality peaked in February and March and were higher the second year when compared with the first.     

Spawning behavior of the pygmy leatherjacket Brachaluteres jacksonianus (Monacanthidae) in southeastern Australia

Spawning behavior of the pygmy leatherjacket Brachaluteres jacksonianus was studied in southeastern Australia. Prespawning females thrust the snout repeatedly into spawning substrata, while males exhibited courtship display and nuzzled the female’s face. The females and males touched their abdomens in pairs and released gametes. No parental egg care was observed after spawning. Fertilized eggs measured 0.74 mm in diameter, and each clutch contained 50–191 eggs. The eggs were attached to filamentous red algae, which were found on the surface of the seagrass and other substrata. Reproductive ecology of B. jacksonianus is similar to that of congeneric Brachaluteres ulvarum, although selection of spawning substrata differed in two species.Supplementary material to this paper is available in electronic format at http://dx.doi.org/10.1007/s10228-005-0272-8     

Sex allocation in a simultaneous hermaphrodite, the zebra goby Lythrypnus zebra: insights gained through a comparison with its sympatric congener, Lythrypnus dalli

Synopsis The sexual pattern of the zebra goby Lythrypnus zebra, is an apparent exception to sex allocation theory. Most L. zebra are simultaneous hermaphrodites (i.e., have active female and male gonadal tissue), yet it appears they do not reproduce as males and females simultaneously. Understanding the maintenance of simultaneous hermaphroditism in L. zebra could expand sex allocation theory. In this study, I used a comparison with the blue-banded goby, Lythrypnus dalli, a sympatric congener with a qualitatively similar sexual pattern, to investigate the role of male spawning rate, body size and sexual flexibility in determining the sexual pattern of L. zebra and to isolate differences between the species that might explain their differing sexual patterns. Using field measurements of male nesting success, I found no differences between the species in the body size of nesting males suggesting that large size is associated with successful male reproduction in both species. In addition, nesting males spawned at approximately three times the rate of females in both species; thus, reproduction via male function can be equally advantageous relative to adopting the female role. However, the nest longevity of L. zebra males was shorter than that of L. dalli males, suggesting reproduction via male function may be less reliable in L. zebra. Finally, under laboratory conditions, L. zebra females tended to prefer large mates, and L. zebra were able to re-allocate in both directions, exhibiting a greater capacity to switch than L. dalli. Given these results, I suggest that switching between the sexes plays a greater role in maintaining simultaneous hermaphroditism in L. zebra than L. dalli, perhaps because male reproduction is not as consistent in L. zebra. Sexual flexibility may be an important factor affecting patterns of sex allocation, generally.     

Reproductive behavior of free-ranging Lemur catta at Beza Mahafaly Special Reserve,Madagascar

Observations of reproductive behavior in free-ranging Lemur catta were carried out during one annual cycle. Variability in the behavior of female ringtailed lemurs during parturition appears to be mainly a function of the female's parity and thus her experience. Females within a troop show estrous asynchrony and characteristically mate with more than one male. Females also exhibit proceptive behavior toward and mate with some males from other troops and with transferring males. The potential for a male to monopolize mating opportunities during a female's short estrous period is therefore limited. Male mating strategies in ringtailed lemurs can be seen as adaptations to female mate choice during a highly restricted breeding season. In this species the dominance hierarchy does not break down with regard to the order of mating. The highest ranking male (central male) mates first and shows precopulatory guarding and longer postejaculatory guarding, which may increase his chances of siring the offspring. Subsequent mating partners have developed various counterstrategies to mitigate mating order effects.