Female resistance and male force: context and patterns of copulation in the New Zealand stitchbird Notiomystis cincta

The New Zealand stitchbird or hihi Notiomystis cincta is unique in that it has two distinct mating positions, in addition to the male standing on the female's back, as is seen in all other birds, it also copulates face‐to‐face. In this study, 43 male stitchbirds first attracted a female to their territory and supplemented their within‐pair matings by intruding into other territories and attempting forced copulations – often resulting in high levels of female harassment. I recorded the temporal variation of both attempted and successful copulations relative to the female's fertile period in order to understand the function of copulation variation in this species. Each of 105 observed copulations were classified according to whether they were: (1) within‐pair or extra‐pair, (2) forced or unforced, and (3) face‐to‐face or standing. Two copulation categories ‘within‐pair unforced standing’ (50%) and ‘extra‐pair forced face‐to‐face’ (27%) accounted for the majority of all observed copulations, and this supported the previous categorisation of face‐to‐face copulation being forced by extra‐pair males in this species. However, in 10% of copulations the female conceded to copulate with an extra‐pair male in a standing position while displaying only subtle signs of resistance, thus, female stitchbirds may show convenience polyandry when the costs of resistance are high. The peak in copulation frequency centred on two days prior to the laying of the first egg and occurred within a period of six days before and seven days after the first egg was laid. Unsuccessful extra‐pair forced copulation attempts closely followed the distribution of all copulations. Male age and morphometrics did not predict whether a female would resist or accept extra‐pair copulation attempts, and female resistance did not appear to depend on male quality. Despite the current trend focussing on female fitness benefits associated with EPCs, it appears that male stitchbirds gain EPCs through forced copulation and females gain no apparent benefit.     

The pelagic ecology of the Grey and Red-necked Phalaropes Phalaropus fulicarius and P. lobatus in the Bay of Fundy, eastern Canada

The outer Bay of Fundy, eastern Canada, is an important feeding area for migrant Grey and Red-necked Phalaropes Phalaropus fulicarius and P. lobatus in late summer. The birds feed on copepod-sized zooplankton, brought to the surface by the passage of strong tidal streams over shallow, rocky 'ledges', and concentrated there in upwelling and convergence 'streaks'. Feeding phalaropes are significantly more abundant in the 'streaks' than in adjacent 'contro' areas. Red-necked Phalaropes are the more common species; they prefer the New Brunswick shore, where copepod biomass is high and the species-community is dominated by large, Stage VI-V Calanus finmarchicus . Grey Phalaropes occur mainly on the Nova Scotian side of Fundy, where the copepod biomass, and the dominant species, are smaller. The feeding habits of both species, and their non-breeding distributions, are reviewed. It is concluded that upwellings, convergences, and other oceanographic phenomena that concentrate zooplankton at the surface are the principal factors influencing the pelagic ecology of phalaropes.     

Mate switching and copulation behaviour in King Penguins

Extra‐pair paternity (EPP) in monogamous birds may result from either extra‐pair copulations (EPCs) or mate switching. In this study of King Penguins in South Georgia, we observed no EPCs at all, an effect of very efficient mate guarding. Onshore males fast and need not divert attention to foraging or to defending nest or territory, as this species has neither. However, we found that mate switching was common. On average 38% (range: 29%–56%; three years pooled) of the birds established pair bonds with at least one initial partner before switching to the partner they bred with (i.e. the “pair mate”). Of the observed copulations of 44 studied females, 22% were with initial partners and 78% with the pair mate. This and the high proportion of mate switching suggest that roughly 10% of the females could have received sperm from males other than the pair mate. The average copulation frequency was 0.026 h^?1, resulting in an estimated 8.2 copulations per clutch (which consists of one egg). That more copulations than necessary for fertilisation occur suggests that males try to protect paternity by sperm competition, and that this is a result of the potential for EPP due to mate switching in King Penguins. All observed copulations except one took place between days 13 and 5, with the peak 7.5 days prior to egg‐laying. The birds found their pair mates (often not the same as in the previous year) on average about 10 days before egg‐laying, and always established themselves at the outskirts of the colony about 8 days before egg‐laying. Thus, most copulations occurred around the time the birds joined the colony. We suggest that it is adaptive to obtain a breeding spot early, because the colony will grow and pairs joining later will protect the offspring. Additionally, we suggest that early copulation outside the colony is adaptive because of the risk of failing to fertilise the egg when copulating among aggressive neighbours inside the dense colony. Based on these two arguments we suggest a “safe place hypothesis” to explain the early copulation peak in King Penguins.     

Sperm Competition and Copulation Intervals of the White Spoonbill (Platalea leucorodia,Aves, Threskiornithidae)

Some aspects of sperm competition were studied in the white spoonbill (Platalea leucorodia) breeding in Doñana National Park (SW Spain). Shorter pair copulation intervals occurred during the prelaying period, when females were subjected to a relatively high frequency of extra-pair copulations. Pair copulation intervals with an intermediate extra-pair copulation by the male mate were longer than those without extra-pair copulation. This result indicates that males need a time of recovery between copulations before they can perform another. Extra-pair copulations by the females did not affect the length of intervals between pair copulations. There were no differences between the lengths of the intervals between an extra-pair copulation by the female and the following pair copulation for cases in which the male mate detected an intruder male attempting copulation with his mate and those in which the intruder remained undetected. However, the correlations obtained between copulatory intervals for detected and undetected cases suggest a copulatory response by their mates, although affected by the required recovery time between copulations by the males. Finally, since extra-pair copulations mainly occurred while male mates were collecting nest material, they engaged in this activity shortly after pair copulations, probably to avoid a last-male advantage under the sperm competition pressure.     

Comparison of behavioral sequence of copulation between chimpanzees and bonobos

We compared sex differences in behaviors leading to copulation of chimpanzees (Pan troglodytes) in the Kalinzu Forest, Uganda with those of bonobos (Pan paniscus) at Wamba, D.R. Congo, using the same definition. Female chimpanzees were more likely to initiate copulation than female bonobos. While most of copulations (96%) were initiated by males in bonobos, among chimpanzees only 63% of copulations were initiated by males. Female bonobos initiated an interaction leading to copulation when males approached them within a short distance. On the other hand, both male and female chimpanzees initiated behavior at a longer distance. Higher proceptivity and a higher copulation rate during the maximal swelling period of female chimpanzees might suggest that they gain greater benefits from a high frequency of copulations than do female bonobos.     

Mating system and nesting biology of the Red-necked Phalarope Phalaropus lobatus: what constrains polyandry?

In this five-year study of Red-necked Phalaropes on the coast of Hudson Bay, Canada, only 8% (5/59) of females were polyandrous. Seventy-five percent were monogamous, and 17% did not obtain mates. All males obtained mates readily. To understand the low incidence of polyandry in this sex-role reversed, non-territorial mating system, aspects of the pair bond, nesting biology, and size and plumage dimorphism were examined.
Pair bonds lasted an average of 11 days and ended as soon as the clutch was completed. Polyandrous females initiated their second nests only seven days later. Since in most years nest initiation for the population as a whole spanned three weeks, there was ample time for multiple clutches. Energetic constraints may have played a limited role in constraining polyandry, but many females that had previously laid clutches courted incubating males or harassed pairs.
The major factor limiting polyandry appeared to be availability of mates. Because phalaropes generally breed in habitats that are less defensible than those of other role-reversed shorebirds, they are less able to monopolize males in populations with equal sex ratios. Sexual size dimorphism is correspondingly smaller than in territorial role-reversed shorebirds, although plumage variation between and within sexes in this study was strong (and invariant with age). Thus, despite the phalaropes' role-reversal, the evidence indicates that monogamy may predominate, with sexual selection driven by the ability of females to produce second clutches opportunistically when receptive males are available.     

Female mate preferences and subsequent resistance to copulation in the mallard

In the majority of socially monogamous bird species, femalessolicit or accept copulations from males other than their partner.Females may gain direct benefits from extrapair males, suchas greater access to resources, or indirect genetic benefitsthat will influence the future success of their offspring. However,one group of birds appears to be the exception to this generalrule; in the wildfowl (Anseriformes), all extrapair copulationsappear to be resisted by females. It has been suggested thatresistance behavior may be a strategy to allow females a greaterchoice of mates, either at the precopulatory level (to promotechoice of copulation partner) and/or the postcopulatory level(to promote multiple mating to increase their choice of sperm).This paper examines the function of female resistance behaviorin one of the dabbling ducks, the mallard (Anas platyrhynchos).Observations on a marked population of wild mallard and experimentswith captive birds found that although females showed a strongpreference for particular males that are the first to molt intotheir breeding plumage, male attractiveness did not influencefemale responses to pair or extrapair copulation attempts. Femaleresistance decreased the likelihood that copulation attemptswould end in successful insemination. The findings did not supportthe hypothesis that females resist copulations to promote femalechoice and the reasons why waterfowl may benefit from avoidingall extrapair copulations are discussed.     

True deception during extra‐pair courtship feeding: cheating whiskered tern Chlidonias hybrida females perform better

Males of many bird species feed their mates during the pre‐incubation period. The food provisioned by males during these courtship feedings (CFs) represents the key source of energy for the female during egg formation. Non‐pair males may trade food for extra pair copulations (EPC) with females during extra pair courtship feeding (EPCF), while females may trade copulations for food with non‐pair males to obtain additional resources. Because EPCs can be costly to the females, they are expected to behave in ways that will deceive non‐pair males to obtain additional resources at no cost to themselves. We investigated EPCFs in whiskered terns Chlidonias hybrida breeding in food‐rich conditions, on carp ponds in southern Poland. Almost all CFs (n = 2751) took place during the female's fertile period and peaked just before clutch initiation. 10% of all CFs were performed by non‐pair males. Females tried to obtain food from the non‐pair male during 39% of EPCFs, by swindling (the female solicits a non‐social male for copulation and tries to swindle food with no cloacal contact) or by snatching (the female tries to take the gift without engaging in copulation). In the remaining 61% of EPCFs, females did not react or chased the visiting male away. The probability of a female's obtaining food during EPCF was much higher (0.69, 95% CI: 0.47–0.85) if she swindled rather than snatched (0.08, 95% CI: 0.02–0.22). Only 0.7% of EPCFs were followed by EPCs. The high availability of food in the study area allows males to perform frequent EPCFs, despite the very low probability of obtaining EPCs. This is the first time that ‘true deception’ during EPCFs has been reported in birds: swindling females obtain food from non‐pair males at no immediate detectable cost to themselves.     

Pair copulation frequency correlates with female reproductive performance in Tree Sparrows Passer montanus

In socially monogamous birds, males benefit from frequent copulations with their partner if this increases their paternity in the brood. By soliciting and accepting copulation attempts, female birds can control pair copulation frequencies. Engaging in copulations can be costly by taking up time and by increasing the risks of predation and pathogen transmission. Current hypotheses propose that high pair-copulation frequencies may compensate for these costs by providing females with direct benefits that can result in higher reproductive success. In this study, I examine in a Tree Sparrow population the prediction that high pair-copulation frequencies are associated with better female reproductive performance. The results show that clutch sizes laid were positively correlated with the pair-copulation frequency. The length of the incubation period was negatively correlated and, as expected, the number of fledglings was positively correlated with pair-copulation frequency. The results are consistent with current hypotheses for the evolution of frequent pair copulations and provide one of the few evidences for a positive relation between pair-copulation frequency and female reproductive performance. This study also suggests that within-pair copulation frequency could be an early expression of the pairs' reproductive ability and might signal their phenotypic quality.     

Sneaky Monkeys: An Audience Effect of Male Rhesus Macaques (Macaca mulatta) on Sexual Behavior

Males and females have different sexual interests and subsequently may show conflicting sexual strategies. While dominant males try to monopolize females, promiscuity benefits females and subordinate males. One way to escape monopolization by dominant males is to copulate in their absence. We tested this inhibitory effect of males on the sexual behavior of their group members in captive group‐living Rhesus macaques. Copulations between females and nonalpha males almost exclusively took place when the alpha male was out of sight. Furthermore, the inhibiting effect was not unique for the alpha male. An upcoming nonalpha male also inhibited copulations of its group members, and three other nonalpha males inhibited female copulation solicitations. Females adjusted their behavior to the presence of bystander males, as they initiated and accepted initiations more often in absence than in presence of bystander males. Although not significant, in males, a similar pattern was found. The observed reduction in mating behavior in presence of bystander males is in accordance with an “audience effect,” in which the behavior is modulated in relation to the presence or absence of third parties. This audience effect may serve as an important mechanism to reduce (aggressive) interruptions of subordinate male copulations.     

Female sexual receptivity and male copula guarding during prolonged copulations in the damselflyIschnura senegalensis (Odonata:Coenagrionidae)

Laboratory experiments were conducted to clarify the relationship between female sexual receptivity and male copula guarding inI. senegalensis, a species that copulates for several hours. In insectaries, most copulations were initiated early in the morning, and terminated relatively synchronously between 11 00 and 13 00. Females refused males with wing-flutter display and oviposited alone in the afternoon regardless of copulation events of that morning. Females could sexually receive males only in the morning. Males copulated for several hours until 12 00 after which females could oviposit. To determine whether copulations that last for hours function as male copula guarding or only of sperm displacement, emerged males were kept at various densities and permitted to copulate with virgin and mated females in insectaries. Both with virgin and mated females, “social” (not solitary; 2–4 males / insectary) males initiated copulations early in the morning and always terminated at around 12 00. However, both with virgin and mated females, solitary (one male / insectary) males terminated copulations in the morning. In both cases, duration of copulations did not significantly differ for virgin females and mated females. Therefore, long (several hour) copulation is more likely to function as male copula guarding than as sperm displacement, and duration of copulations is predicted to be shortened when male density is very low.     

Why Do Female Birds Reject Copulations from their Mates?

Female birds frequently reject copulations from their mates, suggesting a conflict between the sexes. This study analyses behavioural data of socially monogamous razorbills, Alca torda, to examine whether females rejected their mates because of conflicts over fertilization or the pair bond. Among pairs, females rejected 9–70 % of their mates’ copulation attempts and prevented their mates from completing 42–100 % of successful copulations. Copulations terminated by females were half the duration of those terminated by males, and females terminated fewer first copulations than subsequent ones on the same day. These findings indicate that females were motivated to copulate less frequently and for shorter durations than their mates. The sperm competition hypothesis predicts that females reject their mates to increase the probability of being fertilized by extra-pair males. This hypothesis was not supported because females rejected extra-pair males similarly to their mates. The female-mate-guarding hypothesis predicts that females guard their pair bond by copulating frequently with their mates, thereby depriving the males of time and energy to copulate with and form bonds with other females. This prediction was consistent with a significant negative correlation between the percentage of copulation attempts that females accepted from their mates, and the number of extra-pair copulations that their mates attempted. However, this correlation was not caused by a trade-off of males copulating with their mates instead of attempting extra-pair copulation because males attempted most extra-pair copulations on days when their mates were absent. A new hypothesis is proposed, namely, that females reject their mates to test the male's commitment to provide essential parental contributions after egg-laying. The ‘testing-of-the-bond’ hypothesis is consistent with the findings but requires testing.     

Male Competition over Access to Females in a Spider with Last-Male Sperm Precedence

Agonistic behaviour between male cellar spiders (Pholcus phalangioides) was investigated to test whether (1) size difference determines which male achieves access to the female, (2) males are able to monopolize access to the female until egg laying and whether (3) female resource value increases before egg laying because of last‐male sperm precedence. We further investigated whether (4) there is variation in time and energy spent on courtship and copulation depending on the degree of sperm competition, i.e. with or without rival present. In three experimental settings we introduced two males of either different or similar sizes, or a single male to a female. The mating units were constantly video‐observed until the females produced their first egg sac. Experience, ownership and female resource value in terms of body size was controlled. Our results show that larger males achieve almost exclusive access to females. Size symmetrical settings resulted in increased fighting activity and duration but dominance did not influence mating success. If copulations were disturbed by the rival male, copulations were terminated earlier in symmetrical settings compared with asymmetrical settings. In 94.8% of trials only one copulation took place, suggesting that the copulating male successfully monopolized access to the female. Males confronted with a rival copulated longer but courted significantly shorter than lone males. Although the last male to copulate sires 88% of the offspring in P. phalangioides, neither fighting nor courtship activity increased before the female laid a batch of eggs. This suggests that males have no indication of the timing of oviposition.     

Responses of territorial and floater male Red‐winged Blackbirds to models of receptive females

Red‐winged Blackbirds (RWBL; Agelaius phoeniceus) have a polygynous mating system and, because territorial males commonly have harems of two to five females, some second‐year (SY) and after‐second‐year (ASY) males do not establish nesting territories, but become floaters. Previous studies have revealed high rates of extra‐pair copulations in this species and that sexually mature male floaters and territory owners do not differ in size, testosterone levels, or reproductive capability, suggesting that floaters may occasionally gain paternity. During May and June 2008, we observed the behavioral responses of floater males to taxidermic mounts (models) of female RWBL placed in a precopulatory position. Floaters intruded into territories during 46% of model presentations, with 20% of intrusions by ASY floaters and 80% by SY floaters. During intrusions, ASY floaters attempted to copulate with models 93% of the time compared to 80% for SY floaters. Copulations were successful during 30% of attempts by ASY males and 25% of attempts by SY floaters. The frequency of intrusions by ASY and SY floaters, attempted copulations by SY floaters, and successful copulations by ASY floaters increased as territorial males spent more time off their territories. Responses of floater males toward models in our study suggest that floater male RWBL attempt to exploit available breeding opportunities. The lack of evidence for extrapair young (EPY) fathered by floater male RWBL in previous studies, combined with our results indicating that the presence of territorial males limits floater intrusions, copulation attempts, and successful copulations, suggests that the reproductive success of floater males is limited in part by the aggressive behavior of territorial males.     

Rhesus macaque copulation calls: Re-evaluating the “honest signal” hypothesis

Male rhesus macaques sometimes give loud calls while thrusting or dismounting during multi-mount copulations.Hauser (1993) has proposed that these calls (1) impose a cost (increased risk of aggression) on calling males, and (2) increase callers' copulation frequencies, supporting the hypothesis that calls function as honest signals (handicaps) that females use to evaluate male quality during mate choice. This hypothesis was re-examined using data collected at Cayo Santiago, Puerto Rico on 40 focal females and their 56 observed copulatory partners. Although attacks by males against copulating pairs were frequent, they were usually directed only against the female of the pair. Males that called were no more likely than silent males to suffer male escalated attack during or immediately following mount series. Male-female dyads in which the male called during copulation were significantly more likely than non-calling dyads to complete the most copulations observed for any given female. Males that called at least once were significantly more likely than non-calling males to complete at least one copulation with a peri-ovulatory female. A log-linear model revealed that male rank and calling were both associated with likelihood of experiencing at least one peri-ovulatory copulation. However, calling was not associated with reception of demonstrated female mate choice behaviors. Controlling for dominance rank, callers did not experience more female proximity maintenance than non-callers. Nor were callers' hip-grasps refused less frequently than non-callers' hip-grasps. These results cast doubt on the hypothesis that rhesus macaque copulation calls are costly, honest indicators of intrinsic male quality. A contrasting alternative hypothesis, that a male's copulatory calls advertise relative immunity from attacks against his copulatory partners, was not supported either. Thus, the function of rhesus macaque copulatory calls remains unknown. The unusually high rate of copulations amongHauser's (1993) subjects may explain the discrepancy in results, but it is unclear how high copulation rates would increase the cost of copulatory calling to males.     

Pre-oviposition mate-guarding and mating behaviour of Argia vivida (Odonata: Coenagrionidae)

Abstract. 1. At Halcyon Hotsprings, British Columbia, Canada, male and female Argia vivida Hagen encountered to mate in two different ways.
2. In the morning (before 12.30 hours solar time), males basked at sunspots in the forest and darted out at passing females, attempting to take them in tandem (the first method of encounter).
3. If a male was successful, the pair engaged in a 31.3±4.8 min copulation followed by an hour of tandem flight before beginning oviposition.
4. As the day progressed, unmated males moved slowly toward the water and arrived at the water at about the same time as the earliest ovipositing pairs (1131±27.5 min solar time).
5. Males retained their grasp on their mates during oviposition (contact-guarding) but since some tandems separated during oviposition, non-tandem males at the water could capture recently released, gravid females (the second method of encounter).
6. The new pairs performed a brief copulation (10.2±3.38 min) and began ovipositing immediately thereafter.
7. Some females that avoided recapture attempted to oviposit unguarded.
8. We believe the long duration of morning copulations and period of tandem constitute a male strategy, which we call 'pre-oviposition guarding', to guard females until it is warm enough at the oviposition site for the females to begin ovipositing.
9. Separation of tandems during oviposition may be initiated by either member of the pair and we suggest that one benefit to a female of leaving a guarding mate is increased efficiency of oviposition when the intensity of male harassment is low.
10. The mating system of A. vivida thus comprises a series of complementary male and female mating behaviours.     

Mating behaviors of the proboscis monkey (Nasalis larvatus)

The mating behaviors of the proboscis monkey were observed in a riverine forest along a tributary of the Kinabatangan River, Sabah, Malaysia, for a period of 30 months. Solicitation for copulation was initiated frequently by males and occasionally by females. Most copulations involved only one mount; however, some multiple-mount copulations were observed and a maximum of six mounts per copulation were recorded. The mean duration of mounts was about 27 sec. Nonsexual mounts (female-female, female-juvenile/infant, juvenile-juvenile, and juvenile-infant) were also observed. Female-female mounts occurred shortly after failed solicitations toward males were observed. Harassment by juveniles and/or infants was observed during copulation; however, these harassments apparently did not interfere with copulation. Sexual swelling was evident in 77.4% of copulating females, with copulating subadult females showing the most distinct swelling.     

Was tun Buchfinken (Fringilla coelebs) zur Brutzeit au?erhalb ihrer Reviere?

Male and female Chaffinches used small territories and much more extended home ranges. Trips outside their territories were devoted to foraging and other behaviour including extra pair display and copulations, whereas intra pair copulations and some vocalizations were observed exclusively inside the territories. The territory function is discussed.     

Flatworms flatten to size up each other

When copulations are costly, fertilization is reciprocal and fecundity is positively related to size, hermaphrodites are expected to favour large partners, leading to size-assortative mating. Size-related mate choice has, however, never been observed in hermaphrodites. In the flatworm Dugesia gonocephala copulations cost time and are reciprocal, and size is a positive predictor of female fecundity. Every copulation is preceded by a phase in which one partner glides on top of the other and both spread out and flatten, suggesting that partners assess each other''s size. A total of 124 copulating pairs collected on four different dates, proved that mating is size-assorted in the field. In experiments with groups, more copulations took place between equally sized individuals than were expected when matings were random. In experiments with pairs, partners of different size exhibited twice as many mating attempts for the first copulation than did partners of the same size. We conclude that D. gonocephala employs a unique kind of pre-copulatory ''flattening'' behaviour as a mechanism to signal as well as to assess relative size. This does not only confirm that hermaphrodites can mate assortatively when certain assumptions are met, it also proves that even lower invertebrates can show active mate choice.     

Geolocator tagging reveals Pacific migration of Red‐necked Phalarope Phalaropus lobatus breeding in Scotland

The migration route of Red‐necked Phalarope populations breeding on North Atlantic islands has been subject to considerable speculation. Geolocator tags were fitted to nine Red‐necked Phalaropes breeding in northern Scotland to assess whether they migrated to Palaearctic or Nearctic wintering grounds. Of four birds known to return, two had retained their tags, of which one was recaptured. This male Phalarope left Shetland on 1 August 2012 and crossed the Atlantic Ocean to the Labrador Sea off eastern Canada in 6 days, then moved south to reach Florida during September, crossed the Gulf of Mexico into the Pacific Ocean and reached an area between the Galapagos Islands and the South American coast by mid‐October, where it remained until the end of April, returning by a similar route until the tag battery failed as the bird was crossing the Atlantic Ocean. The total migration of 22 000 km is approximately 60% longer than the previously assumed route to the western part of the Arabian Sea, and this first evidence of migration of a European breeding bird to the Pacific Ocean also helps to indicate the possible migratory route of the large autumn movements of Red‐necked Phalaropes down the east coast of North America.