Search PubMed⌕ Search

SEARCH · Search PubMed

Results for “COURTSHIP”

Search indexed PubMed citations on genomics, clinical trials, systematic reviews and public health. Explore titles, authors and supplied subject terms, then open the PubMed record.

Quote a phrase for an exact phrase match. Source license links do not imply unrestricted reuse.

At least 523 records · Page 29Linked to original sources
PubMed2005

The effects of genotype, age, and social environment on male ornamentation, mating behavior, and attractiveness.

The traits thought to advertise genetic quality are often highly susceptible to environmental variation and prone to change with age. These factors may either undermine or reinforce the potential for advertisement traits to signal quality depending on the magnitude of age-dependent expression, environmental variation, and genotype-age and genotype-environment interaction. Measurements of the magnitude of these effects are thus a necessary step toward assessing the implications of age dependence and environmental variability for the evolution of signals of quality. We conducted a longitudinal study of male guppies (Poecilia reticulata) from 22 full-sibling families. Each fish was assigned at maturity to one of three treatments in order to manipulate his allocation of resources to reproduction: a control in which the male was kept alone, a courtship-only treatment in which he could see and court a female across a clear partition, and a mating treatment in which he interacted freely with a female. We measured each male's size, ornamental color patterns, courtship, attractiveness to females, and mating success at three ages. Size was influenced by treatment and age-treatment interactions, indicating that courtship and mating may impose costs on growth. Tail size and color patterns were influenced by age but not by treatment, suggesting fixed age-dependent trajectories in these advertisement traits. By contrast, display rate and attempted sneak copulation rate differed among treatments but not among ages, suggesting greater plasticity of these behavioral traits. As a result of the different patterns of variation in ornamentation and behavior, male attractiveness and mating success responded to male age, treatment, and the interaction between age and treatment. Neither age nor treatment obscured the presence of genetic variation, and the genetic relationship between male ornamentation and attractiveness remained the same among treatments. Our findings suggest that neither age-dependent variation nor environmentally induced variation in reproductive effort is likely to undermine the reliability of male signaling.

Age Factors↗
PubMed1977

Factors influencing ovulation in the Mexican axolotl as revealed by induced spawnings.

Axolotl females may be induced to spawn by injecting them with small doses of FSH prior to mating. Data on the different categories of spawnings obtained by this procedure have established the following points. (1) Spawnings of injected females completely without fertile eggs are small (173 eggs average). Courtship occurred in some of the matings (spermatophores found) but not in all. Spawnings of injected but uninseminated females re-mated with a second (or third) male, not becoming inseminated, but all having some eggs fertilized by contact with spermatophores (showing courtship) are likewise small (219). It can be concluded that courtship of the injected female, even by two successive males, does not stimulate ovulation of additional eggs. (2) Insemination of injected females induced additional ovulation more or less frequently, depending upon whether the female was inseminated by the first, or a subsequent male with whom she was mated. Insemination by the first male, within 15 hours after the female had been injected with FSH, resulted in stimulation of ovulation in 36 of 54 spawnings (67%). The 36 spawnings averaged 593 eggs per spawning. In contrast, insemination by a second or third male, more than 15 hours after the female was injected with FSH, resulted in stimulation of additional ovulation in only 4 of 21 spawnings (19%). (3) Survival time of spermatozoa in inseminated females has been established. Considerable numbers of spermatozoa capable of fertilizing eggs survive for about one week. A few survive for 11 to 12 days, but none was found after 14 days.

Ambystoma↗
PubMed2006

Lesion-induced neurogenesis in the hypothalamus is involved in behavioral recovery in adult ring doves.

Although neurogenesis in the brain of adult vertebrates is region dependent, lesion induces generation of new neurons in non-neurogenic brain regions. These findings raise the question of the role of new neurons in brain repair and functional recovery. We addressed this question by applying previous observations that electrolytic lesion induced neurogenesis in the ventromedial nucleus (VMN) of the hypothalamus in adult ring doves. Such lesions disrupted the male's courtship behavior, which could be reinstated after rehabilitation with a female. We investigated whether lesion-induced newborn neurons in the VMN facilitate the recovery of courtship behavior in the lesioned birds. We conducted systematic observations of cytological, morphological, and neuroanatomical changes in the lesioned VMN, and concurrently we monitored behavioral changes. Using a multitude of specific cell markers, we found a well-circumscribed cellular zone that proliferated actively. This highly proliferative zone initially appeared along the periphery of the lesion site, where cells had high levels of expression of neuronal, glial, and neurovascular markers. As newborn neurons matured at the lesion site, the necrosis gradually decreased, whereas a downsized proliferative zone relocated to a region ventral to the VMN. Some of the mature neurons were found to project to the midbrain vocal nuclei. Restoration of these projection neurons coincided with the recovery of courtship vocalization. Finally, we found that a social factor, that is, when the male doves were cohoused with a mate, facilitated neurogenesis and behavioral recovery. These results suggest that lesion-induced neurogenesis contributes to behavioral recovery in adult animals.

Aging↗
PubMed1997

Vocal behaviour of the frog Eleutherodactylus antillensison the British Virgin Islands

The vocal repertoire of male Eleutherodactylus and the communicative significance of two calls (trill and courtship call) were investigated under natural conditions on the British Virgin Islands. Interspersed with two-note advertisement calls, males uttered trill calls, which typically consisted of an introductory note followed by five to 13 rapidly repeated notes. The introductory note and the repeated notes were within the range of the dominant frequencies of the first and second note of the advertisement call, respectively, but their mean frequencies were 5-8% higher. The temporal and spectral characteristics of the trill and advertisement calls were similar on three islands (Tortola, Virgin Gorda, Guana), except that the number of repeated notes in the trill was higher on Guana. Body size and air temperature had no detectable effects on the call characteristics. In response to experimental playback of trill calls (either synthesized or constructed from actual calls), males produced more trills in the period following the stimulus than during the baseline. During the playback itself, five or 22 males produced an unusually high number of trills that alternated with the stimulus trills. These responses suggest that the trill is an aggressive call used in the defence of a calling territory. In response to the playback of courtship calls (multi-note calls that males uttered only in the presence of a female and that differed from the trill call mainly in the dominant frequency of the repeated notes), 11 of 28 males left their perch sites and nine approached the speaker. In contrast, males never left their perches during the baseline or the trill stimulus. These responses suggest that males may exploit the courtship calls of rivals to obtain mates through interference.

Journal Article↗
PubMed1997

Behaviourally mediated sexual selection: characteristics of successful male black grouse

Recent studies of lekking animals suggest that a suite of characters may be favoured by sexual selection. Examples of such traits are high survival, increased androgen levels, territory features and morphological characters including exaggerated morphological ornaments. Here we argue that such selection is often mediated by behavioural differences and we use results from a field study of lekking black grouse, Tetrao tetrixas an example of our argument. In the absence of females, males display a range of stereotyped behaviours including vocalizations, visual displays and fighting. When females attend the lek, the behavioural repertoire of the males becomes more limited. Time budget analyses reveal that in the vast majority of cases, males in the presence of females perform only three types of behaviour: a vocal display called rookooing; a courtship behaviour called circling; and fighting. This suggests that female preference could be based on male courtship and fighting behaviour while females visit individual male territories and that displays not used in the presence of females could be ruled out as important for female preference. Sexual selection in black grouse, however, is a complex interplay between male-male competition and female choice and therefore female preference is not the only determinant of male success. Successful males were more often involved in fights in the absence of females on the lek and defended territories that were larger than expected. Therefore, we hypothesize that differences in male fighting ability result in some males occupying relatively large territories at the centre of the leks. Relatively large territories seem to be needed for successful courtship. Therefore behavioural differences mediate differences in male copulation success both through female preference and male-male competition.

Journal Article↗
PubMed1998

Tibial coloration, fluctuating asymmetry and female choice behaviour in the damselfly Platycypha caligata.

The territorial damselfly Platycypha caligata (Odontata: Chlorocyphidae) has a courtship behaviour where males wave the white anterior surface of all six laterally enlarged tibiae at females. I experimentally altered this white tibial surface using black paint to determine the effect on male behaviour of a 25% reduction in area, or an increase in asymmetry between the left and right side of the body. I collected behavioural data on courtship, mating and fighting for males already holding territories. Neither a reduction in the area nor an increase in asymmetry of tibial whiteness affected male mating rate, courtship rate or fighting behaviour. These manipulations also had no significant effect on the daily presence at the study site. Males whose tibial whiteness was experimentally removed also showed no decrease in mating rate or change in fighting behaviour while territorial. The complete removal of tibial whiteness did, however, lead to a significant reduction in daily presence, possibly due to a reduced ability to acquire or hold a territory. There was no relationship between natural levels of asymmetry in tibial whiteness and mating rate, courting rate or fighting behaviour for males with territories. However, the natural area of tibial whiteness was significantly positively related to both mean male mating rate and copulation duration for territorial males. This result suggests that a phenotypic correlate of area of tibial whiteness, probably body size, is sexually selected through female choice among males that already hold territories.Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed1998

Female fifteen-spined sticklebacks prefer better fathers.

We studied how male fifteen-spined sticklebacks, Spinachia spinachia, vary in paternal competence, whether males advertise their competence and whether females prefer better fathers. In this species the male alone provides care for the offspring through nest building, fanning, cleaning and protecting the eggs. We found no female preference for larger males. Instead, females preferred males that during the subsequent paternal phase fanned their nests in shorter fanning bouts. Such males enjoyed a significantly higher hatching success because they fanned more often than males with longer fanning bouts. Males that fanned for short bouts during the paternal phase were also able to increase their fin beat rate. Frequent fanning and high fin beat rates may improve the flow of oxygen to the eggs. Beat rate may be a condition-dependent trait, because males that lost more weight were unable to increase their fin beat rate. During courtship, males perform behaviours such as displacement fanning and body shaking. Females preferred males showing more frequent body shakes during courtship. Body shake frequency correlated positively with hatching success, and negatively with mean fanning bout duration during the paternal phase. The results indicate that female S. spinachia show a preference for direct benefits in terms of better paternal care, and that males may signal this ability to females by shaking their body during courtship. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed1998

Coevolution of elaborated male display traits in the spotted bowerbird: an experimental test of the threat reduction hypothesis.

Elaborated male sexual displays commonly involve multiple elements that may differ greatly among closely related species, but there have been few studies of the causes of this divergence. Male spotted bowerbirds, Chlamydera maculata, have unusually intense, aggressive courtship displays and highly divergent bowers. Male and female courtship positions differ from related species in that males court females separated by a modified see-through bower wall. Here we experimentally tested hypotheses that could explain the unique features of spotted bowerbird display relative to other Chlamydera species. Our results support the threat reduction hypothesis, which suggests that accessory traits evolve because they mitigate the threat associated with intense, aggressive male displays that are most effective in causing females to become sexually receptive. In spotted bowerbird males, the highly modified and unique see-through bower walls allow females to view intense displays while reducing threat that would otherwise be associated with these same displays. We found that (1) females preferred males with the most intense displays, (2) males and females at experimentally manipulated bowers consistently moved to courtship positions so that the standing bower wall separated them and (3) males reduced display intensity when not separated from the female by an intact bower wall. Comparisons with other Chlamydera species suggest that bower architecture coevolves with other display elements to allow maximally effective male displays. Such coordinated adaptive changes of display elements suggest an alternative to the runaway divergence of arbitrary traits as the cause of rapid divergence in multifaceted male display traits among closely related species. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed1995

Endocrine basis of the reproductive pattern of the Gentoo penguin (Pygoscelis papua): winter breeding and extended laying period in northern populations.

Changes in plasma LH, prolactin, testosterone, estradiol, and progesterone were investigated throughout moult and reproduction in free-living male and female Gentoo penguins (Pygoscelis papua) at Crozet Island (46 degrees S, 51 degrees E), where this species is able to relay after a reproductive failure. In both sexes, LH, prolactin, and steroid hormones, remained at basal levels during the moult. LH level was highest at the time of arrival at the colony for breeding and, although it decreased after courtship, it did not drop at basal value by incubation and first chick brooding period. Prolactin peaked for both chick brooding periods; replacement clutch was associated with an increased secretion of LH, whereas high prolactin levels were maintained. Testosterone, in male, and estradiol, in female, peaked during courtship I and chick brooding II; progesterone, in female, peaked during courtship I and II. These hormonal patterns are consistent with those observed in passerine species which are also able to relay after a reproductive failure. Winter breeding observed at Crozet Island might reflect the extreme adaptive capacity of Gentoo penguin species.

Adaptation, Biological↗
PubMed2001

Behavioral and hormonal effects of exogenous vasotocin and corticosterone in the green treefrog.

Vasotocin (AVT) promotes courtship in a wide range of vertebrates. However, this effect is not independent of steroid hormones. For example, androgens may work in concert with AVT and corticosterone (CORT) may work to oppose AVT action. In frogs, AVT promotes calling, and in some species, CORT inhibits calling. In addition, androgens are known to modulate AVT in the brain, and CORT may depress androgen secretion. Previous work in amphibians has suggested that AVT promotes courtship by overcoming a CORT-mediated stress response. Possible behavioral and hormonal interactions among AVT, CORT, and androgens were investigated in wild, free-living green treefrogs (Hyla cinerea). Saline, AVT, CORT, or a combination of AVT and CORT were administered to calling males, and several measures of spontaneous calling were evaluated for 1.5 h following injection. Plasma testosterone, dihydrotestosterone, and CORT were also measured. Saline-injected males had low CORT levels, and AVT and CORT injection elevated plasma CORT levels. AVT increased the likelihood of calling, but, in males who did call, AVT did not influence latency to call or how often they were observed calling. Very few saline-injected males resumed calling after injection, and therefore a CORT effect was only detectable in AVT-injected males. CORT inhibited calling in AVT-injected males only at the highest dose of CORT (40 microg); lower levels of CORT were unsuccessful at inhibiting AVT-induced calling. AVT appeared to have a specific effect on calling motivation. Further, the data suggest that disinhibition of a CORT response is not the primary mechanism by which AVT increases calling. In addition, CORT injection reduced endogenous androgen levels. Finally, endogenous androgens were negatively correlated with latency to begin calling, suggesting that they may have a positive effect on calling. These data indicate that AVT has positive effects on calling but provide only weak evidence that CORT inhibits courtship in this species.

Animals↗
PubMed1995

Hormonal correlates of breeding status, nest construction, and parental care in multiple-brooded northern mockingbirds, Mimus polyglottos.

Blood samples were taken from free-living male and female mockingbirds to determine correlations between plasma steroids and breeding behavior. Plasma levels of testosterone (T), estradiol (E2), progesterone (P), and corticosterone (B) were assessed during pre-breeding courtship, while males built nests, while females incubated, while parents fed young, and while males built new nests as pairs continued to care for dependent fledglings. Samples were also taken from unmated males holding spring territories. In males, T and P fluctuated with breeding phase, and in all groups, male T levels exceeded basal values measured in autumn. But, when mated males built nests in the absence of young, T levels were higher than in all other conditions, though only for early broods. In females too, T and P levels changed with breeding phase; T levels rose during courtship and while males built first nests and were intermediate during nest construction for later broods. Male and female P levels rose during courtship and when males built nests with young present. While feeding young in the nest, males showed significantly higher corticosterone levels than females, though in females, both P and B levels increased during incubation for replacement broods. These findings strengthen an earlier finding that T may activate male nest construction, and they suggest that breeding behavior may be related to T in females as well. Progesterone data raise the possibility that P too, may be linked to breeding behavior in male and female mockingbirds. Sex differences in corticosterone levels during the nestling period may reveal subtle costs associated with male mockingbirds' highly aggressive defense against predators.

Animals↗
PubMed1998

Effects of 17alpha-methyltestosterone on sexually dimorphic characters in the weakly discharging electric fish, Brienomyrus niger (Günther, 1866) (Mormyridae): electric organ discharge, ventral body wall indentation, and anal-Fin ray bone expansion.

Adult males of African weakly discharging electric fish (family: Mormyridae) are distinguished from juveniles and adult females by a dorsally directed indentation of the posterior ventral body wall and by massive bone expansion of the bases of a select number of anal-fin rays. These sexually dimorphic structures seem to facilitate the anal-fin reflex that is displayed during courtship when the male envelopes its anal fin around the female's to form a common spawning pouch. Expanded bone could provide additional surface for muscle attachment and thus assist in part with the courtship sequence. Based on the fact that the expression of the male sexually dimorphic electric organ discharge (EOD) is under androgen control, and that the female EOD can be masculinized through testosterone administration, we hypothesized that androgens should also drive anal-fin ray bone expansion in male mormyrids and equally effect male-like changes in treated juveniles and adult females. Exogenous androgen treatment (17alpha-methyltestosterone) of adult female Brienomyrus niger resulted in a male-like EOD, and male-typical structural transformations (body wall indentation and anal-fin ray bone expansion). Some of these changes were immediate and receded following hormone withdrawal (EOD), while others developed more slowly and were apparently permanent (indentation and bone formation). 17alpha-Methyltestosterone administration affected only those targets in females that are normally involved in the male's reproductive behavior, i.e., its courtship signal (EOD) and two morphological features (body-wall indentation and bone expansion). Rays of the dorsal or caudal fins were never affected.

Animals↗
PubMed1999

Lesions to the medial preoptic area affect singing in the male European starling (Sturnus vulgaris).

The aromatization of testosterone (T) in the medial preoptic nucleus (POM) is known to regulate male courtship and sexual behaviors expressed prior to, and in anticipation of, copulation. Singing in male European starlings is used to attract mates prior to physical sexual contact, suggesting that the POM might be involved. The present study was performed to examine the effects of lesions targeting the POM on singing and courtship behavior in reproductively active male starlings. A significant decrease in song output and the gathering of green nest materials was observed in males with lesions to the POM compared to males with damage to brain areas outside of the POM. Lesions did not affect a male's tendency to remain near a female or to occupy a nestbox, suggesting that the effects of POM lesions were specific to courtship behaviors. Behavioral differences were not related to testis mass or volume, and GnRH immunoreactivity was observed within the hypothalamus and median eminence for each male, suggesting that the effects of POM lesions were related specifically to POM involvement in song expression rather than to a disruption of the GnRH axis. These results suggest a general role for the POM in the expression of behaviors related to sexual arousal or anticipation, including song.

Analysis of Variance↗
PubMed2002

Seasonal and sexual dimorphisms in the green anole forebrain.

During the breeding season, male green anole lizards extend a throat fan (dewlap) in courtship. This behavior is facilitated by testosterone (T). Females extend a much smaller dewlap less often, even with the same dose of T. During the nonbreeding season when T is low, dewlap extension is reduced. To determine if parallels exist between structure and function, we investigated neuron soma size and density in the preoptic area (POA) and ventromedial nucleus of the amygdala (AMY), which are involved in the display behavior, in breeding and nonbreeding males and females. Cells from breeding animals were larger than cells from nonbreeding animals, but they were not sexually dimorphic. No significant effects existed in neuron density. This experiment indicates that portions of the anole forebrain important for sexual behavior are plastic and might be influenced by seasonal changes in steroid hormones. To investigate whether T can reverse the seasonal difference in soma size in both sexes, gonadectomized nonbreeding anoles were implanted with an empty or T propionate-filled capsule; animals were also tested for male-typical courtship behavior. Males and females treated with T had higher rates of dewlap extension, but across treatment groups these rates were greater in males. Neuron soma size in the POA and AMY was larger in males than females, but no effects of treatment were detected. Taken together, the results indicate that T can stimulate behavior in the nonbreeding season and suggest that a dissociation exists between the regulation of the courtship display and soma size of relevant brain regions.

Aggression↗
PubMed2002

Context-dependent effects of castration and testosterone treatment on song in male European starlings.

Most seasonally breeding songbirds display dramatic seasonal fluctuations in plasma testosterone (T) levels and mate attraction behaviors, including song. However, males of some songbird species, such as the European starling (Sturnus vulgaris), continue to sing at high levels after the breeding season, when T levels are basal. In male starlings song during the breeding season functions mainly to attract mates, whereas song during the nonbreeding season appears unrelated to reproduction. This suggests that song produced in a context unrelated to female courtship, unlike song directed toward females, is not regulated by plasma T. In captive males housed in large outdoor aviaries we explored the relationship between plasma T and song produced during the breeding season within and outside a courtship context. This was achieved by determining the effects of castration and subsequent T treatment on song and mate attraction behaviors in both the presence and the absence of a female. Compared to intact males, castrated males did not show reduced song activity in the absence of a female for at least 6 months after the operation, strongly suggesting that the expression of noncourtship song is not regulated by plasma T. Likewise, we found that experimentally elevating T levels in castrated males did not affect noncourtship song rates. However, control castrated males receiving empty implants tended to show reduced noncourtship song rates after implantation. This may have been due to a suppressive effect caused by the presence of the T-implanted castrated males in the same aviary. In contrast, courtship singing was clearly controlled by plasma T: it was abolished by castration and restored by subsequent T replacement when males were housed both individually and in a group situation. High plasma levels of T also appeared necessary for the activation of three other behavioral traits critical for mate attraction, namely, nesthole occupancy, spending time (singing) in a nesthole, and carrying green nesting material into a nesthole.

Animals↗
PubMed2001

Phylogeny of Limia (Teleostei: Poeciliidae) based on NADH dehydrogenase subunit 2 sequences.

Complete sequences for the mitochondrial gene NADH Dehydrogenase 2 (ND2) and partial sequences for the tRNA-Met and tRNA-Trp genes were obtained for 11 populations of the poeciliid fish genus (or subgenus) Limia, including species from Hispaniola, Cuba, Jamaica, and Grand Cayman Islands. Additional sequences for Limia (2 species), Pamphorichthys (3), Poecilia (16), and Xiphophorus (1), all from the tribe Poeciliini, were extracted from GenBank, as was a sequence for Heterandria formosa, from the tribe Heterandriini. Phylogenetic analyses included parsimony, distance methods, and maximum-likelihood. Several Limia species groups that had been proposed based on morphological data were evaluated. The versicolor species group was strongly supported, as was the close relationship between the Cuban Limia vittata and the Grand Cayman Limia caymanensis. However, the proposed vittata species group was not upheld by the ND2 data. The phylogenetic position of Limia melanogaster, from Jamaica, was either sister to the versicolor species group or basal to all other included Limia species. Limia was found to be monophyletic; however, Limia species from the island of Hispaniola were not monophyletic. There was little support for any proposed sister group to Limia. The phylogeny was used to reconsider a previous comparative study of poeciliid courtship behavior and sexual dimorphism. The data indicated that there may have been two independent appearances of courtship display behavior in Limia; considering the tribe Poeciliini as a whole, there is evidence for as many as five appearances of display (including two within Limia), or a complex pattern of gains and losses of such behavior. The application of phylogenetic information to the comparative study did not refute the previously hypothesized correlation between the presence of a courtship display and the presence of sexual dimorphism in poeciliid fish.

Animals↗
PubMed1986

Acoustic communication in an electric fish, Pollimyrus isidori (Mormyridae).

It has been known since von Frisch's work in the 1930's that mormyrid electric fishes are quite sensitive to sound. We now describe a repertoire of natural sounds produced by the mormyrid, Pollimyrus isidori, during breeding and aggression; reception of communication sounds is probably a major function for mormyrid audition. In aquaria, Pollimyrus isidori produce 'grunts', 'moans', 'growls', 'pops' and 'hoots' at various phases during nesting, courtship, and territory defense. All five sounds are produced primarily at night. Territorial males produce grunts, moans and growls during courtship. Vocalizing is stimulated by the presence of a gravid female on the male's territory and decreases with the onset of spawning. Hoots and pops are given during agonistic behavior. Grunts are bursts of acoustic pulses, stereotyped for an individual, with the potential as individual signatures. The electric organ is silent during grunts and moans and is discharged at a reduced rate during growls. The courtship and spawning of Pollimyrus isidori is described.

Agonistic Behavior↗
PubMed1977

Portions of the central nervous system controlling reproductive behavior in Drosophila melanogaster.

Drosophila melanogaster sex mosaics were tested in their courtship interactions with females and then with males. The distribution of genetically male and female tissues in each mosaic was determined with respect to an external cuticle marker and an internal enzyme marker. Performance of malelike courtship was correlated with the genotype of various tissues, with special attention being paid to the genotypes of head and thoracic ganglia. Male tissue in the left or right dorsal brain is necessary and nearly always sufficient to trigger early courtship actions--following of females and wing extension at them--but male tissue in both the dorsal brain and thoracic ganglia is necessary for attempted copulation to occur. Female tissue on or in the abdomen is nearly always necessary and sufficient for a mosaic to be courted by a male.

Animals↗