Paternity analysis of two male mating tactics in the fiddler crab, Uca mjoebergi

The fiddler crab Uca mjoebergi mates both underground in male-defended burrows and on the surface near female-defended burrows. The reproductive tract of Uca species facilitates last-male precedence, suggesting that males that do not guard-mated females are likely to gain very little paternity if the female re-mates with another male. Here, we test the reproductive success of burrow and surface matings using paternity analysis. We found that 100?% of the females that mated in burrows extruded a clutch of eggs. Furthermore, we show conclusively, for the first time in a fiddler crab species, that last-male sperm precedence results in the majority of the female’s eggs being fertilised by the burrow-mated male. In contrast, surface matings resulted in significantly fewer females extruding eggs (5.6?%). Paternity analysis also revealed that more than half of the clutches from burrow-mated females showed low levels of extra-pair paternity from previous matings. Although multiple matings appear common in U. mjoebergi, burrow-mated males that guard females are guaranteed a successful mating with extremely high rates of assured paternity. Surface matings therefore appear to be an opportunistic tactic that may increase male reproductive success in a highly competitive environment.     

Multiple paternity, sperm storage, and reproductive success of female and male painted turtles (Chrysemys picta) in nature

When females receive no direct benefits from multiple matings, concurrent multiple paternity is often explained by indirect genetic benefits to offspring. To examine such possibilities, we analyzed genetic paternity for 1,272 hatchlings, representing 227 clutches, from a nesting population of painted turtles (Chrysemys picta) on the Mississippi River. Goals were to quantify the incidence and distribution of concurrent multiple paternity across clutches, examine temporal patterns of sperm storage by females, and deduce the extent to which indirect benefits result from polyandrous female behaviors. Blood samples from adult males also allowed us to genetically identify the sires of surveyed clutches and to assess phenotypic variation associated with male fitness. From the genetic data, female and male reproductive success were deduced and then interpreted together with field data to evaluate possible effects of female mating behaviors and sire identity on offspring fitness. We document that more than 30% of the clutches were likely fathered by multiple males, and that presence of multiple paternity was positively correlated with clutch size. Furthermore, the data indicate that the second male to mate typically had high paternity precedence over the first.     

Female reproductive qualities affect male painted turtle (Chrysemys picta marginata) reproductive success

Male reproductive success (RS) in polygamous species with minimal social systems is often determined by the number of mates. However, because male RS is translated through females, the number of offspring sired can also be influenced by female qualities. Empirically sufficient data to document how tradeoffs between mate number and quality influence male RS are seldom available for long-lived, iteroparous species. We combined long-term life history data (1983–2006) on the E. S. George Reserve (ESGR, MI, USA) with parentage data from 155 clutches of 59 female painted turtles (Chrysemys picta marginata) of varying reproductive frequencies (2003–2006) to determine the relative contribution of female numbers and qualities on male RS. One previously documented trait of female painted turtles that can have substantial influences on male RS is repeat paternity through the use of stored sperm to fertilize over 95 % of within-year clutches. In addition, our study found that second-clutch producing female painted turtles on the ESGR have higher among-year reproductive frequencies than do first-clutch only females. Multiple paternity was detected in 14.1 % of clutches (min-max?=?6.1–30.0 % annually), and the number of mates of both sexes was low annually (males 1.0; females 1.2) and over 4 years (males 1.1; females 1.7). Among successful males, RS varied substantially (1–32 offspring) and was strongly influenced by the combination of female reproductive frequency and repeat paternity (>38 % among years), but not mate number. Low mate number for both sexes was unexpected in a species without complex mating behaviors or parental care.     

Sex-specific reproductive behaviours and paternity in free-ranging Barbary macaques (Macaca sylvanus)

In a wide variety of species, male reproductive success is determined by contest for access to females. Among multi-male primate groups, however, factors in addition to male competitive ability may also influence paternity outcome, although their exact nature and force is still largely unclear. Here, we have investigated in a group of free-ranging Barbary macaques whether paternity is determined on the pre- or postcopulatory level and how male competitive ability and female direct mate choice during the female fertile phase are related to male reproductive success. Behavioural observations were combined with faecal hormone analysis for timing of the fertile phase (13 cycles, 8 females) and genetic paternity analysis (n = 12). During the fertile phase, complete monopolisation of females did not occur. Females were consorted for only 49% of observation time, and all females had ejaculatory copulations with several males. Thus, in all cases, paternity was determined on the postcopulatory level. More than 80% of infants were sired by high-ranking males, and this reproductive skew was related to both, male competitive ability and female direct mate choice as high-ranking males spent more time in consort with females than low-ranking males, and females solicited copulations mainly from dominant males. As most ejaculatory copulations were female-initiated, female direct mate choice appeared to have the highest impact on male reproductive success. However, female preference was not directly translated into paternity, as fathers were not preferred over non-fathers in terms of solicitation, consortship and mating behaviour. Collectively, our data show that in the Barbary macaque, both sexes significantly influence male mating success, but that sperm of several males generally compete within the female reproductive tract and that therefore paternity is determined by mechanisms operating at the postcopulatory level.     

Females reward courtship by competing males in a cannibalistic spider

Despite widespread recognition that intersexual interactions shape reproductive strategies, studies of male competition do not typically include effects imposed by females. In cannibalistic redback spiders, escalated fighting between rival suitors is predicted, as males are unlikely to mate with more than one female, and strong first-male sperm precedence favours mating with virgins. In staged competitions for matings between size mismatched rivals, smaller males adopted an alternative sneaking strategy. However, despite initial agonistic interactions, larger males did not pursue or incapacitate smaller males. When inter-male competition occurred, females struck at males frequently, although strikes were rarely seen when males courted in the absence of a rival. After minimal fighting, larger males engaged in significant courtship (3 h) rather than killing inferior rivals. Prolonged courtship was favoured by female behaviour, as males that attempted rapid copulation (smaller, sneaking males) were cannibalised before mating was completed. This premature cannibalism significantly decreases paternity in redback spiders. Thus, significant features of male competitive behaviour (i.e. prolonged courtship by larger males) may be predicted with consideration of the female’s response to male reproductive strategies. Although the effect of females may be more subtle in systems without the extreme reversed size-dimorphism of redbacks, these results suggest that female interests should be explicitly considered when studying inter-male interactions.     

Determinants of paternity success in the spider Pholcus phalangioides (Pholcidae: Araneae): the role of male and female mating behaviour

In double mating experiments, we examined whether and to what extent various male and female behavioural traits influence the course of mating and fertilization success in the cellar spider. In males, we focussed on pre-copulatory behaviour and on the rhythmic twisting movements that the male performs with his pedipalps during copulation. In females, we investigated remating decisions and the effect of female termination of copulation. Second males fertilized a high proportion of the eggs (P ₂: median 89%) despite much shorter second matings, with high variation in relative paternity success. The number of pedipalp movements (PPMs) of either male was a better predictor of paternity than copulation duration. Our results suggest that in second matings, PPMs help to remove sperm from previous males, whereas in first matings a high number of PPMs enhances fertilization success, either due to numerical sperm competition or cryptic female choice. Furthermore, we found a negative male age effect on paternity in second matings, implying that age-related deterioration of spermatozoa may promote variation in fertilization success. Female receptivity decreased significantly in second matings; only 70% of the females remated. Females that accepted a second copulation were found to terminate these much earlier and with higher probability than first matings. This suggests that the intensity of conflict between the sexes is higher in second matings. Increased intensity of sexual conflict may be responsible for stronger selection on male traits, as pre-copulatory behaviour and age only affected male copulatory performance and paternity in second matings. Electronic Publication     

Male remating and female fitness in the wolf spider Pardosa astrigera: the role of male mating history

Although the effects of male mating history on female reproductive output and longevity have been studied in insects, few such studies have been carried out in spiders. In a mating system in which females are monandrous while males are polygynous, females may incur the risk by mating with successful males that have experienced consecutive matings and suffer from the possible depletion of sperm and/or associated ejaculates. Here, we examine the effects of male mating history on male courtship and copulation duration, female reproductive fitness, and female adult longevity of the wolf spider, Pardosa astrigera. Results indicated that male mating frequency had little effect on their subsequent copulation success, and of 35 males tested, about half of the males were able to copulate with five virgin females successively at an interval of 24 h. Male mating history had little effect on their courtship duration. However, male mating history significantly affected male copulation duration, female adult longevity, and reproductive output. Males that mated more frequently copulated longer and more likely failed to cause their mates to produce a clutch, although there was no significant difference in the number of eggs laid and the number of eggs hatched regardless of the first clutch or the second one. Multiple mating of male P. astrigera resulted in significant reduction in female adult longevity. Our results indicate that monandrous females mating with multiple-mated males may incur substantial fitness costs.     

Female choice in a promiscuous wild guinea pig,the yellow-toothed cavy (<Emphasis Type="Italic">Galea musteloides</Emphasis>)

Promiscuity is traditionally considered to increase only male reproductive success but, more recently, female benefits are also assumed to be the driving force for promiscuous mating. The yellow-toothed cavy (Galea musteloides) is characterised by an extremely high degree of multiple paternity (>80%), and females have greater offspring viability after mating multiple males. However, so far it is not clear whether or not it is the female's decision that leads to mating with more than one male. To elucidate the female's role in bringing about promiscuity, female yellow-toothed cavies were given the choice between four different males each, in a mate-choice apparatus that simultaneously prevented monopolisation and harassment of the females by the males. In 10 of the 12 choice tests, mating occurred. Nine of these ten females actively sought copulations with more than one male, and their mating behaviour was displayed in a way that might have favoured the mixing of sperm. At the same time, they significantly preferred heavier and more frequently courting males. These results show that female yellow-toothed cavies are actively involved in mating with more than one male. Thus, the present study is the first to show that in a species in which females produce more viable offspring as a consequence of polyandrous mating, these females are indeed motivated to actively bring about promiscuity. Nevertheless, females are selective in the choice of their mates. Thus, both sperm competition and direct female choice seem to be important for the increased offspring viability due to promiscuous mating.     

Elucidating the function of ejaculate expulsion and consumption after copulation by female Euxesta bilimeki

Postcopulatory processes can influence male reproductive success in several animal species. Females can use different mechanisms to bias male paternity after copulation. One of such mechanisms consists in expelling all or part of the ejaculate after copulation. Euxesta bilimeki is an Ulidiid fly whose females not only frequently expel ejaculates after mating but also consume the ejaculate after expulsion. In order to understand the significance of these behaviors we examined video recordings of courtship, copulatory, and postcopulatory behaviours. The presence of sperm in female storage organs was confirmed after mating with males of different sizes and was correlated with duration of courtship, copulation, and the period from the end of copulation to ejaculate expulsion. The effect of ejaculate consumption on female fitness (fecundity and longevity) was compared among females held under different dietary treatments: a rich diet consisting of protein, sugar and water, an intermediate diet composed of sugar and water, a poor diet of only water and females that were completely deprived of food and water. All of the observed females expelled ejaculates after mating. The probability of storing sperm in the two spermathecae and the ventral receptacle was correlated with interactions between the duration of all behaviours examined and male size. Except for starved females, who lived longer when allowed to consume ejaculates, ejaculate consumption had no effect on fitness. Results suggest that females can bias sperm storage according to male mating effort, while the consumed ejaculate had some nutritional value only evident when females were completely starved.     

Inbreeding avoidance in a poeciliid fish (Heterandria formosa)

Polyandry is hypothesized to give females an opportunity to avoid inbreeding through postcopulatory selection mechanisms if precopulatory inbreeding avoidance is not possible, for example, because of forced matings. Here, we report a postcopulatory, prefertilization, inbreeding avoidance mechanism in the least killifish, Heterandria formosa, a species in which males can force matings. Females had 50% less sperm in their oviducts and ovarian cavities after mating with a sibling compared to a mating with a nonsibling male. Neither sex showed inbreeding avoidance in a dichotomous precopulatory mate choice test or during mating trials. Females in this species invest substantially in each offspring after fertilization (matrotrophy), whereas males invest little more than sperm. Based on theory, females should therefore be more likely than males to avoid inbreeding in this species. We suggest that females do this by reducing the amount of sibling sperm in their reproductive system. However, the possibility that males invested more sperm to nonsiblings could not be ruled out. In a fertilization success experiment, the first male to mate with a female sired all the offspring in most cases, even if it was a sibling. However, large females were more likely to carry offspring of multiple males. Possibly female sperm storage sites were filled by the first male, and only large females had space for the second male's sperm.     

Influence of alternate reproductive tactics and pre- and postcopulatory sexual selection on paternity and offspring performance in a lizard

Sexual selection theory predicts different optima for multiple mating in males and females. We used mating experiments and genetic paternity testing to disentangle pre- and postcopulatory mechanisms of sexual selection and alternate reproductive tactics in the highly promiscuous lizard Eulamprus heatwolei. Both sexes mated multiply: 30–60 % of clutches were sired by two to four fathers, depending on the experiment. Larger males sired more offspring when we allowed male contest competition: 52 % of large males but only 14 % of small males sired at least one offspring. In the absence of male contest competition, females mated promiscuously and there was no large male advantage: 80 % of large males and 90 % of small males sired at least one offspring, and there was no evidence for last-male precedence. Multiple mating did not yield obvious direct or indirect benefits to females. E. heatwolei represents a complex system in which males attempt to improve their fertility success by limiting rivals from access to females and through adopting alternate reproductive tactics. Conversely, females exhibit no obvious precopulatory mate choice but may influence fitness through postcopulatory means by either promoting sperm competition or through cryptic female choice. Our results support the hypothesis that female multiple mating in nonavian reptiles is best explained by the combined effect of mate encounter frequency and high benefits to males but low costs to females.     

Positive association between social and extra-pair mating in a polygynous songbird, the dickcissel (Spiza americana)

In polygynous species, males appear to gain additional offspring by pairing with multiple females simultaneously. However, this may not be true if some females copulate outside of the social pair bond. Polygynous males could experience lower paternity because of trade-offs among gaining multiple social mates, guarding fertility with these mates, and pursuing extra-pair matings. Alternatively, polygynous males could simultaneously gain extra social mates and have high paternity, either because of female preferences or because of male competitive attributes. We tested four predictions stemming from these hypotheses in a facultatively polygynous songbird, the dickcissel (Spiza americana). Unlike most previous studies, we found that males with higher social mating success (harem size) also tended to have higher within-pair paternity and that the number of extra-pair young a male sired increased significantly with his social mating success. Females that paired with mated males were not more likely to produce extra-pair young. In contrast, extra-pair paternity was significantly lower in the nests of females whose nesting activity overlapped that of another female on the same territory. This pattern of mating was robust to differences in breeding density. Indeed, breeding density had no effect on either extra-pair mating or on the association between polygyny and paternity. Finally, nest survival increased with harem size. This result, combined with the positive association between polygyny and paternity, contributed to significantly higher realized reproductive success by polygynous male dickcissels.     

Factors influencing paternity in multiply mated female red-sided garter snakes and the persistent use of sperm stored over winter

In some species, sperm is stored within the female reproductive tract for months to years, and yet remains viable to fertilize eggs and produce offspring. Female red-sided garter snakes store sperm for over 7 months of winter dormancy. In previous work, we demonstrated that these stored sperm account for an average of 25 % paternity of a litter when the female mates with a male at spring emergence. Here, we tested whether last-male sperm precedence was prevalent when a female mates with two males during the spring. On average, paternity was shared equally among the first (P₁ proportion of paternity of the first male to mate) and second males (P₂) to mate in the spring, and stored sperm (P_ss), but the variance in paternity was high. Thus, last male sperm precedence may diminish when a female has more than two mates. Male size did not affect paternity, but, as the interval between matings increased, P₁ increased at the expense of P_ss. Interestingly, as the second spring male’s copulation duration increased, P₁ also increased at the expense of P₂. This result suggests that female influence over sperm and/or copulatory plug transfer during matings may also affect which male fathers her offspring in response to coercive matings as we assisted females to mate for their second mating. Finally, all females were spring “virgins”; consequently, sperm stored from autumn matings (and/or previous spring matings) remain competitive even when faced with two rivals in sperm competition and is likely the driver of the evolution of sperm longevity.     

Female benefit,male risk: Polyandry in the true armyworm Pseudaletia unipuncta

In Lepidoptera polyandry is common and females may increase their lifetime reproductive output through repeated matings if they acquire essential resources from male ejaculates. However, the paternity of males mating with previously-mated females is far from assured unless sperm precedence is absolute. In this study on the polyandrous armyworm, Pseudaletia unipuncta, we used two strains of male (the black-eyed wild type and a red-eyed homozygous, recessive mutant), mated with red-eyed females, to determine (i) whether male investment has any impact on female reproductive output, and (ii) if females do benefit from multiple matings, to what extent males fertilize the eggs to which they contributed. Multiple mating resulted in a significant increase in both the fecundity and longevity of females. However, the degree of sperm precedence (those eggs fertilized by the second male) varied from 0–100%, but was not affected by either male size or age, or by the duration of copulation. In cases where sperm precedence was <50% (x = 12%) females produced significantly more eggs (1384 versus 940) prior to the second mating than females where sperm precedence was >50% (x = 89%), indicating that the quality of the first mating influenced the fertilization success of the female's second mate.     

High degree of multiple paternity in the viviparous Shiner Perch, <Emphasis Type="Italic">Cymatogaster aggregata</Emphasis>, a fish with long-term female sperm storage

The Shiner Perch (Cymatogaster aggregata Gibbons) exhibits a viviparous reproductive mode and long-term female sperm storage, two biological features that may predispose this fish species for both intense sperm competition and frequent multiple paternity within broods. To test these hypotheses, we used polymorphic microsatellite markers to identify sires and quantify paternal contributions to the progeny arrays of 27 pregnant females from a natural population of C. aggregata. The number of sires per brood ranged from one to eight (mean 4.6), typically with skewed distributions of fertilization success by the fathers but no correlation between sire number and brood size. The extraordinarily high incidences of multiple paternity in this species probably are due in part to high rates of mate encounter, but selection pressures related to the avoidance of maternal–fetal incompatibility may further have promoted the evolution of polyandrous mating behaviors in this female-pregnant species. Our genetic data are consistent with the hypothesis that viviparity, long-term sperm storage, and extreme polyandry are interrelated reproductive phenomena that should promote the evolution of post-copulatory sperm competition and/or cryptic female choice in these fishes.     

Estimating paternity with spatial behaviour and DNA fingerprinting in the striped plateau lizard, Sceloporus virgatus (Phrynosomatidae)

The association between spatial proximity and paternity was studied in a population of the striped plateau lizard, Sceloporus virgatus. The relationship between estimated mating success and male phenotypic traits was examined for a sample of 55 males. DNA samples were obtained from 13 female-offspring families. The males with the closest spatial proximity to each female were tested as possible sires within each family. Fingerprinting with two multilocus hypervariable minisatellite probes revealed a strong correspondence between male-female spatial proximity and actual paternity. Paternity could be assigned for 72 of the 100 hatchlings. Most hatchlings with identifiable sires were attributed to a male with the highest category of spatial proximity to the mother. However, there was a low to moderate level of multiple paternity within clutches, and for some clutches probable sires could not be identified even though the most likely behavioural candidates were tested. Thus, nonterritorial males or other males lacking strong social and spatial relationships with females may achieve some degree of reproductive success. Analysis of mating success revealed that male success increased with body size, up to a point beyond which larger size conferred no advantage. Received: 7 January 1997 / Accepted after revision: 16 June 1997     

Fitness benefits of multiple mating versus female mate choice in the cellar spider (Pholcus phalangioides)

Recent studies have demonstrated that mating with multiple males can be beneficial for females and her offspring even if males contribute nothing but sperm. This was mainly established for species in which sperm from several males mix in the reproductive tract of the female, thus allowing sperm competition and/or female sperm choice. However, in species with last male sperm precedence, female re-mating decides against the previous male by strongly limiting his reproductive success. We tested the effect of female re-mating behaviour using the cellar spider Pholcus phalangioides, which shows strong last males sperm precedence and moderate levels of polyandry under natural situations. We predicted that females prevented from remating even though they are receptive would show reduced reproductive success compared to females that accept two copulations and females that reject a second male, since the latter two treatments were allowed to behave according to their decisions. However, if the number of matings per se had an effect on oviposition or on offspring performance, double-mated females should perform better compared to both treatments of once-mated females. We measured female fecundity and fertility over a period of 140 days, comparable to the species' natural reproductive peak season. Two thousand one hundred and fifty-two offspring from 67 first egg sacs were reared under two feeding levels. We registered development time and survival, and measured offspring adult size and mass. We found a positive effect of double mating, as in this treatment, oviposition probability was higher compared to the other treatments. Interestingly, adult female offspring of the DM treatment that were raised under low food level had a higher condition index compared to those from FS and RM, but development time, size and mass at adulthood were not affected by mating treatment. Female choice only seemed to affect hatching latency of the offspring. Overall, the main predictor of female reproductive output and success was female body size.     

Determinants of paternity in the milkweed leaf beetle

Summary Males and females of the milkweed leaf beetle mate for up to 2.5 d in the field; copulation is interspersed with mounted courtship behavior and periods of passive riding on the female's back. In 1985, females mated with up to 10 different males in Bridgeport, New York, and matings averaged 0.75±0.04 (SE) d in duration. Sperm utilization patterns are complex in the milkweed leaf beetle. In the laboratory, mating duration, mating order and the duration of the gap between matings affected paternity. The second male's sperm predominated when a female's two consecutive matings were of equal duration (but were long: 45 h each) or when they were intermediate in duration (15 h each) with a 5 day gap between the two matings. The mechanism of sperm predominance appears to be one of time-dependent sperm removal and replacement. Neither relative nor absolute size of males affected the extent to which they superceded the sperm of rivals. Electrophoretic analysis of paternity at 6 polymorphic loci showed that offspring of at least eight out of 11 mating pairs pairs collected in the field at the end of the 1984 breeding season were sired by more than one male. At least seven of the 11 families produced some offspring that were sired by the female's most recent mate in the field and in no case was the collected male completely excluded from paternity. In a greenhouse plot, males climbed off females' backs a mean of 1.1±0.3 h before oviposition. Females did not oviposit with males on their backs and did not oviposit sooner when males were removed than when males were allowed to remain mounted, indicating that males don't keep females from ovipositing until they can achieve sperm predominance. The close concordance between male disembarkment and oviposition suggests that males remain with females beyond the time when they have superceded the sperm of their prior mates, possibly to guard them from rival males. Although the prolonged mating association of the milkweed leaf beetle cannot be separated into discrete copulatory and guarding phases, the results of this study suggest that prolonged mating enables males both to remove and replace the sperm of the females' prior mates with their own sperm and to prevent females from remating prior to oviposition.     

Sperm economy and limitation in spiny lobsters

Sperm limitation, when female fertilisation success is constrained by the supply of sperm, is generally perceived to be an uncommon feature of reproduction in species which directly transfer gametes during copulation. Male size, previous copulations, and the balance of expected reproductive return and future mating opportunity may, however, limit the amount of sperm males transfer to females. We used laboratory experiments where mate size could be manipulated and its consequences on spermatophore size and clutch size determined, to show that in two genera of spiny lobsters (Crustacea: Palinuridae) male reproductive output limits the size of clutches brooded by females. In Panulirus argus from the Florida Keys, we show that while male size affects spermatophore area, males also vary the amount of ejaculate positively with female size. Furthermore, the area of the spermatophore has a greater influence than female size on subsequent clutch weight. In Jasus edwardsii from New Zealand, female size, male size and mate order all affect clutch weight. In both species, clutches fertilised by small males in the laboratory are significantly smaller than clutches fertilised by large males. These results suggest that to ensure they receive sufficient sperm, females should either mate several times prior to oviposition, mate as early as possible in the reproductive season, or choose large, preferably unmated males as partners and thus compete with other females for preferred males. Sperm-limited female fecundity has the potential to limit the egg production of fished populations where large males are typically rare. Received: 18 May 1998 / Received in revised form: 20 November 1998 / Accepted: 30 November 1998     

High degree of paternity loss in a species with alternative reproductive tactics

In many mating systems, males adopt alternative reproductive tactics (ARTs) to maximize reproductive success. In fishes, guarding males often invest more energy into courtship, defense, and paternal care, whereas cuckolding males forego such costs and steal fertilizations by releasing their sperm in the nest of a guarding male. These two tactics have been documented in the plainfin midshipman fish (Porichthys notatus), yet the relative reproductive success of the guarding and cuckolding male tactics remains unknown. In this study, we used microsatellite markers to determine the level of paternity of the guarding type I males. We explored how paternity varied with male phenotype and across the breeding season. Our results revealed the lowest documented levels of paternity in a species with obligate paternal care. Although paternity remained consistently low, it did increase as the breeding season progressed. Male body size did not significantly predict paternity. The low paternity in this species may be explained, in part, by aspects of their reproductive ecology including the duration of parental care period, limited nest availability and competition for nests, as well as the occurrence of nest takeovers. Overall, our findings contribute to the understanding of the ultimate factors underlying ARTs in this species and highlight the importance of investigating reproductive success across the entire breeding season.