Female Calls Research Articles

Female copulation calls vary with male ejaculation in captive olive baboons

Abstract Copulation calls are mating-associated vocalizations that are common in primates, with females vocalizing after copulation in several Old World monkeys and apes. Baboon females typically produce copulation calls that correlate with fertile phase. Calls are, thus, regarded as an upshot of cycle physiology and sexually selected calls. Here, we describe three captive troops of olive baboons wherein, against expectation, females suppressed vocalizing during copulations. Vaginal cytology, together with sexual swelling observations, confirmed that females experienced full receptive cycles. Ovulation did not affect vocal probability during sex, while copulation calls were predicted by male ejaculation just as in other Old World primate species. Results cast doubt on the existence of physiological triggers for baboon copulation calls. Social factors may instead play a larger role. Alterations in social structure (as typically observed in the wild) may be implemented strategically as captive enrichment in order to reveal how females in highly social primates change sexual strategies and, therefore, the use of their copulation calls.

Female cuckoo calls elicit vigilance and escape responses from wild free-range chickens

Mimicry is widespread in the animal kingdom. Parasitic cuckoos can reduce the risk of their offspring being rejected by employing morphological and egg mimicry. Female common cuckoos (Cuculus canorus) can deceive their hosts by mimicking sparrowhawk (Accipiter nisus) calls, thus improving brood parasitism success. However, few experiments have confirmed the hypothesis that female cuckoos can successfully mimic sparrowhawk calls. We used two populations of wild free-range chickens (Gallus gallus domesticus), which are not cuckoo hosts, to study the effects of four different types of bird calls on their anti-predatory behavior. Results showed that both female cuckoo and sparrowhawk calls elicited vigilance and escape responses from chickens, while the chickens did not respond to either male common cuckoo or Oriental turtle dove (Streptopelia orientalis) calls. This suggests that female cuckoo calls can not only deceive their hosts, but also successfully deceive their non-host birds. Our study further confirms the accuracy with which female cuckoos can mimic sparrowhawk calls.

Neurotelemetry Reveals Putative Predictive Activity in HVC during Call-Based Vocal Communications in Zebra Finches.

Premotor predictions facilitate vocal interactions. Here, we study such mechanisms in the forebrain nucleus HVC (proper name), a cortex-like sensorimotor area of songbirds, otherwise known for being essential for singing in zebra finches. To study the role of the HVC in calling interactions between male and female mates, we used wireless telemetric systems for simultaneous measurement of neuronal activity of male zebra finches and vocalizations of males and females that freely interact with each other. In a non-social context, male HVC neurons displayed stereotypic premotor activity in relation to active calling and showed auditory-evoked activity to hearing of played-back female calls. In a social context, HVC neurons displayed auditory-evoked activity to hearing of female calls only if that neuron showed activity preceding the upcoming female calls. We hypothesize that this activity preceding the auditory-evoked activity in the male HVC represents a neural correlate of behavioral anticipation, predictive activity that helps to coordinate vocal communication between social partners.SIGNIFICANCE STATEMENT Most social-living vertebrates produce large numbers of calls per day, and the calls have prominent roles in social interactions. Here, we show neuronal mechanisms that are active during call-based vocal communication of zebra finches, a highly social songbird species. HVC, a forebrain nucleus known for its importance in control of learned vocalizations of songbirds, displays predictive activity that may enable the male to adjust his own calling pattern to produce very fast sequences of male female call exchanges.

Modulation of Reproductive Behavior of Diaphania indica (Lepidoptera: Pyralidae) by Preferred and Non-preferred Host Plants

The cucumber moth, Diaphania indica (Lepidoptera: Pyralidae), is one of the most destructive pests of cucurbitaceous plants all over the world. Previously, we showed that volatile organic compounds (VOCs) from cucumber plants play a crucial role in guiding gravid D. indica females to oviposition sites, whereas watermelon VOCs did not, and females significantly prefer cucumber over watermelon for oviposition. It remained unclear, however, whether preferred and non-preferred host plants also affect moth reproductive behaviors. In the present study we investigated the effect of preferred and non-preferred host plants on female calling, male attraction to female sex pheromone, and mating behavior in D. indica. In the presence of cucumber, virgin females engaged in significantly more calling on the first night than in either the watermelon treatment or the control. Also, we found that the mean time per bout on the first night in the cucumber plant treatment was significantly higher than that of the other two groups. The combination of cucumber foliage with female pheromone significantly increased the attraction and landing of males compared to either watermelon or the control in the wind tunnel. Furthermore, for one-day-old insects mating occurred only in the cucumber treatment. All insects in the cucumber treatment exhibited their first mating on one of the first three nights, but in the watermelon treatment only by night four had all insects mated. We also found that the duration of first mating in cucumber was significantly greater than in either watermelon or control. Overall, our study provides evidence that preferred and non-preferred host plants affect reproductive behaviors in D. indica.

Female-female aggression and male responses to the two colour morphs of female common cuckoos

Female-only colour polymorphism is rare in birds, but occurs in brood parasitic cuckoos (Cuculidae). Obligate brood parasites leave incubation and parental care to other species (hosts), so female-female interactions can play a role in how parasites guard critical resources (host nests) within their laying areas. The plumage of adult female common cuckoos (Cuculus canorus) is either rufous (typically rare) or grey (common), whereas adult male conspecifics are monochromatic (grey). In previous studies, hosts and conspecific males responded with less intensity toward the rare female morph in support of a negative frequency-dependent benefit of female plumage polychromatism. Here, we assessed responses of both conspecific females and males to vocal playbacks of female calls, coupled with one of two 3D models of the different morphs of female cuckoos. At our study population, the rufous female morph was as common as the grey morph; therefore, we predicted similarly high rates of conspecific responses in both treatments. Both female and male cuckoos responded to playbacks acoustically, which demonstrated the primary role of acoustic communication in social interactions amongst cuckoos. Following this, some cuckoos flew closer to the models to inspect them visually. As predicted, no significant differences were detected between the live cuckoos’ responses toward the two colour morphs in this population. We conclude that dichromatism in female cuckoos evolved to serve one or more functions other than conspecific signalling.

Geographic variation in sexual communication in the cotton bollworm, Helicoverpa armigera.

BACKGROUNDGeographic variation in male response to sex pheromone lures has been studied in the field in a number of moth species. However, only a few studies have investigated geographic variation in female calling and sex pheromone under field conditions. For an effective field implementation of sex pheromone lures, it is essential to know the local sex pheromone blend and local timing of sexual communication. We investigated the level and extent of geographic variation in the sexual communication of the important agricultural pest Helicoverpa armigera (Lepidoptera, Noctuidae) in three continents.RESULTSWe found there is no genetic variation in the calling behavior of H. armigera. In the female sex pheromone, we found more between‐population variation than within‐population variation. In male response experiments, we found geographic variation as well. Strikingly, when adding the antagonistic compound Z11‐16:OAc to the pheromone blend of H. armigera, significantly fewer males were caught in Australia and China, but not in Spain. This variation is likely not only due to local environmental conditions, such as photoperiod and temperature, but also to the presence of other closely related species with which communication interference may occur.ConclusionFinding geographic variation in both the female sexual signal and the male response in this pest calls for region‐specific pheromone lures. Our study shows that the analysis of geographic variation in moth female sex pheromones as well as male responses is important for effectively monitoring pest species that occur around the globe. © 2020 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Six molecular patterns leading to hemophilia A phenotype in 18 females from Poland

IntroductionFemale hemophilia is an intriguing rare disorder and few larger reports on its genetic etiology are available. While historically the diagnosis was satisfactorily reached by factor VIII activity assays, the clinical and potentially therapeutic heterogeneity of female hemophilia calls for comprehensive molecular diagnosis in each case. Currently, the genetic investigations are not a part of routine, state-funded, diagnostics in Poland, and thus molecular epidemiological data are missing. AimWe set out to perform a comprehensive genetic analysis of Polish females with hemophilia A. Patients/methodsEighteen females with hemophilia A (including 2 with severe and 5 with moderate hemophilia phenotype) consented for genetic diagnostics. To establish F8 mutations, we used next-generation sequencing of a panel of genes associated with hematological disorders, standard assays for recurrent intragenic F8 inversions and MLPA when deletions were suspected. When appropriate we also used karyotyping, genomic microarrays and X chromosome inactivation assays. ResultsWhile abnormally skewed X-chromosome inactivation combined with a F8 variant on the active allele was, as expected, the most common genetic etiology, a number of other genetic scenarios were unraveled. This included: misdiagnosis (molecular diagnosis of vWd), Turner syndrome, compound heterozygosity and androgen insensitivity syndrome (a phenotypical 46,XY female with a novel androgen receptor gene mutation). We report 3 novel F8 mutations. ConclusionEvery case of female hemophilia warrants full genomic diagnostics, as this may change the diagnosis or reveal broader morbidity than a coagulation disorder (Turner syndrome, androgen insensitivity, or cardiovascular morbidity that we described previously in a SHAM syndrome carrier).

Identification of a female-produced pheromone in a destructive invasive species: Asian longhorn beetle, Anoplophora glabripennis

Male-produced pheromone components have been reported in the Asian longhorn beetle, Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae), but field attraction to these components has been less than overwhelming. Female ALBs were observed rhythmically extending the genitalia in a manner reminiscent of female calling behaviors in other cerambycid species. We thus hypothesized that female ALBs release volatile pheromone while performing this sex-specific behavior. A group of sesquiterpenes, including a major compound α-longipinene and several minor ones α-cubebene, α-ylangene, (−)-α-copaene, α-bergamotene, β-caryophyllene, and α-farnesene, found in genitalia extracts from virgin females elicited male antennal responses. Y-tube olfactometer assays indicated significant attraction of α-longipinene to both sexes in either the presence or absence of host volatiles. This compound was also detected in genitalia extracts from virgin males, though in much lower quantities than in females. Dose–response experiments conducted in the y-tube olfactometer and field both revealed that α-longipinene was attractive at the higher doses, but not at the lower ones. In the field, traps baited with a blend containing α-longipinene, α-cubebene, and β-caryophyllene captured significantly more ALB than solvent controls. The trap catches of α-longipinene combined with either the minor components or host compounds were both greater than those of α-longipinene alone, but the difference was not significant. These results indicate that α-longipinene is a new type of female-produced volatile pheromone in ALB, and the attraction may be synergistically enhanced by several minor components. Sesquiterpenes may play an important role in intraspecies chemical communication of this insect.

Sexual Behavior of the Sugarcane Hairy Borer, Hyponeuma taltula (Lepidoptera: Erebidae): Evidence for a Female-Released Sex Pheromone.

The sugarcane hairy borer, Hyponeuma taltula (Schaus, 1904), attacks the base of sugarcane and causes the death of the apical bud, a symptom known as "dead heart." In recent years, there has been significant infestation in Brazilian sugarcane crops. With the purpose of finding methods for controlling and/or monitoring the pest for the sustainable cultivation of sugarcane, we characterized the mating behavior of H. taltula and confirmed the release of a sex pheromone by this insect. Sexual activity was observed between the second and ninth hour of scotophase, during which female calling caused the onset of male courtship behaviors and increased their frequency. An ethogram was built showing that males perform seven observable steps: (1) wing flapping, (2) walking with wing flapping, (3) flying, (4) flying towards the female, (5) landing near the female and walking around her, (6) attempting copulation, and (7) copulation (mating). Female sex gland extracts also triggered attraction and male courtship behaviors. These results thus confirm that the female abdominal gland produces a sex pheromone and that the presence of this pheromone in the air is needed to initiate the male courtship sequence. The sexual behavior of H. taltula is compatible with control and/or monitoring methods using female sex pheromones.

Black-capped Chickadees (Poecile atricapillus) can identify individual females by their fee-bee songs

Abstract Individual recognition is a social behavior that occurs in many bird species. A bird’s ability to discriminate among familiar and unfamiliar conspecifics is critical to avoid wasting resources such as time and energy during social interactions. Black-capped Chickadees (Poecile atricapillus) are able to discriminate individual female and male chick-a-dee calls, potentially male and female tseet calls, and male fee-bee songs. In the current study, we used an operant discrimination go/no-go paradigm to determine whether female and male chickadees could discriminate between fee-bee songs produced by individual female chickadees as well as test which song component(s) enable this discrimination. Birds trained on natural categories—the songs of different females—learned to respond to rewarded stimuli more quickly than birds trained on random groupings of female songs and were able to transfer this learning to new songs from the same categories. Chickadees were also able to generalize their responding when exposed to the bee note of the fee-bee song of rewarded individuals; they did not generalize to fee notes. Our results provide evidence that Black-capped Chickadees can use female-produced fee-bee songs for individual recognition. However, the acoustic features underlying individual recognition require further investigation.

Sex differences in vocalization are reflected by event-related potential components in the music frog.

Sex differences in vocalization have been commonly found in vocal animals. It remains unclear, however, how animals perceive and discriminate these differences. The amplitudes and latencies of event-related potentials (ERP) components can reflect the auditory processing efficiency and time course. We investigated the neural mechanisms of auditory processing in the Emei music frog (Nidirana daunchina) using an Oddball paradigm with ERP. We recorded and analyzed eletroencephalogram (EEG) signals from the forebrain and midbrain when the subjects listened to white noise (WN) and conspecific sex-specific vocalizations. We found that (1) both amplitudes and latencies of some ERP components evoked by conspecific calls were significantly higher than those by WN, suggesting the music frogs can discriminate conspecific vocalizations from background noise; (2) both amplitudes and latencies of most ERP components evoked by female calls were significantly higher or longer than those by male calls, implying that the ERP components can reflect sex differences in vocalization; and (3) there were significant differences in ERP amplitudes between male and female subjects, suggesting a sexual dimorphism in auditory perception. Together, the present results indicate that the music frog could discriminate conspecific calls from noise, male's calls from female's ones, and sexual dimorphism of auditory perception existed in this species.

Reciprocal Matched Filtering in the Inner Ear of the African Clawed Frog (Xenopus laevis).

Anurans (frogs and toads) are the most vocal amphibians. In most species, only males produce advertisement calls for defending territories and attracting mates. Female vocalizations are the exceptions among frogs, however in the African clawed frog (Xenopus laevis) both males and females produce distinct vocalizations. The matched filter hypothesis predicts a correspondence between peripheral auditory tuning of receivers and properties of species-specific acoustic signals, but few studies have assessed this relationship between the sexes. Measuring hearing sensitivity with a binaural recording of distortion product otoacoustic emissions, we have found that the ears of the males of this species are tuned to the dominant frequency of the female's calls, whereas the ears of the females are tuned close to the dominant frequency of the male's calls. Our findings provide support for the matched filter hypothesis extended to include male-female calling. This unique example of reciprocal matched filtering ensures that males and females communicate effectively in high levels of background noise, each sex being most sensitive to the frequencies of the other sex's calls.

Generation, Coordination, and Evolution of Neural Circuits for Vocal Communication.

In many species, vocal communication is essential for coordinating social behaviors including courtship, mating, parenting, rivalry, and alarm signaling. Effective communication requires accurate production, detection, and classification of signals, as well as selection of socially appropriate responses. Understanding how signals are generated and how acoustic signals are perceived is key to understanding the neurobiology of social behaviors. Here we review our long-standing research program focused on Xenopus, a frog genus which has provided valuable insights into the mechanisms and evolution of vertebrate social behaviors. In Xenopus laevis, vocal signals differ between the sexes, through development, and across the genus, reflecting evolutionary divergence in sensory and motor circuits that can be interrogated mechanistically. Using two ex vivo preparations, the isolated brain and vocal organ, we have identified essential components of the vocal production system: the sexually differentiated larynx at the periphery, and the hindbrain vocal central pattern generator (CPG) centrally, that produce sex- and species-characteristic sound pulse frequencies and temporal patterns, respectively. Within the hindbrain, we have described how intrinsic membrane properties of neurons in the vocal CPG generate species-specific vocal patterns, how vocal nuclei are connected to generate vocal patterns, as well as the roles of neurotransmitters and neuromodulators in activating the circuit. For sensorimotor integration, we identified a key forebrain node that links auditory and vocal production circuits to match socially appropriate vocal responses to acoustic features of male and female calls. The availability of a well supported phylogeny as well as reference genomes from several species now support analysis of the genetic architecture and the evolutionary divergence of neural circuits for vocal communication. Xenopus thus provides a vertebrate model in which to study vocal communication at many levels, from physiology, to behavior, and from development to evolution. As one of the most comprehensively studied phylogenetic groups within vertebrate vocal communication systems, Xenopus provides insights that can inform social communication across phyla.

Long-term Devocalization of Zebra Finches.

Songbirds, such as the zebra finch, are a popular animal model for studying the neural basis of vocal and complex skill learning. Adult male zebra finches produce courtship song toward females (referred to as 'directed song') and recording and analyzing sounds of directed song along with underlying neural activity is important for investigating behavioral and neural mechanisms of song production and learning. However, recording of directed song is easily contaminated by calls that are often as loud as directed songs and frequently produced by a female bird is presented in the same sound-recording chamber to elicit directed song. We developed a new surgical procedure to relatively easily and almost completely devocalize female zebra finches semi-permanently, without affecting other behaviors. This procedure enables researchers to record directed songs with almost no contamination by female calls. The procedure can also be used to devocalize male birds as well and, thus, has great potential for a variety of experimental purposes, such as long-term elimination of auditory feedback during singing in male birds.

Monogamy in a Moment: How do Brief Social Interactions Change Over Time in Pair-Bonded Zebra Finches (Taeniopygia guttata)?

SynopsisResearch on monogamy has largely focused on marked behaviors that are unique to pair bonded partners. However, these marked behaviors represent only a subset of the pair-directed behaviors that partners engage in; the influence of pair bonding on mundane or subtle social interactions among partners remains largely unknown. In this study, we describe the changes that occur during brief social reunions (or greets) over the course of pair bonding in zebra finches. We quantified pair-directed behavior during 5-min reunions from three stages of pair bonding: initial pairing (between 4 and 72 h), early pairing (1–2 weeks), and late pairing (>1 month). These social interactions were operationalized in multiple ways. First, we quantified the overall activity levels (call and movement rates) for both the male and female. Overall, females were more active than males, but for both males and females calling activity was highest at initial pairing. We quantified behavioral coordination between partners in two ways: (1) similarity in call and movement rates between partners and (2) temporal synchrony of calls and movements between partners (via sliding correlation coefficients of time-stamped calls and movements). Overall, there were no effects of pairing stage on behavioral coordination. Finally, we used principal component analyses to disentangle behavioral coordination from the activity levels of the male and female. These results contribute to a growing line of evidence that male and female zebra finches differentially contribute to social dynamics and highlight the influence of pair bonding on the development of social dynamics. Furthermore, our preliminary analyses raise the hypothesis that behavioral coordination during the earliest phases of pairing is modulated by the extent and nature of prior experience. Overall, while behavioral coordination is clearly important for many salient interactions such as duetting, courtship displays, and biparental care, the significance of mundane social interactions for monogamous partnerships remains largely unknown.

The influence of long-term cadmium exposure on the male advertisement call of Xenopus laevis

Cadmium (Cd) is a non-essential environmental endocrine-disrupting compound found in water and a potential threat to aquatic habitats. Cd has been shown to have various short-term effects on aquatic animals; however, evidence for long-term effects of Cd on vocal communications in amphibians is lacking. To better understand the long-term effects of low-dose Cd on acoustic communication in amphibians, male Xenopus laevis individuals were treated with low Cd concentrations (0.1, 1, and 10μg/L) via aqueous exposure for 24months. At the end of the exposure, the acoustic spectrum characteristics of male advertisement calls and male movement behaviors in response to female calls were recorded. The gene and protein expressions of the androgen receptor (AR) were determined using Western blot and RT-PCR. The results showed that long-term Cd treatment affected the spectrogram and formant of the advertisement call. Compared with the control group, 10μg/L Cd significantly decreased the first and second formant frequency, and the fundamental and main frequency, and increased the third formant frequency. One and 10-μg/L Cd treatments significantly reduced the proportion of individuals responding to female calls and prolonged the time of first movement of the male. Long-term Cd treatment induced a downregulation in the AR protein. Treatments of 0.1, 1, and 10μg/L Cd significantly decreased the expression of AR mRNA in the brain. These findings indicate that long-term exposure of Cd has negative effects on advertisement calls in male X. laevis.

Effect of Diallyl Trisulfide on the Reproductive Behavior of the Grain Moth, Sitotroga cerealella (Lepidoptera: Gelechiidae).

The Angoumois grain moth, Sitotroga cerealella (Olivier, 1789) (Lepidoptera: Gelechiidae), is primarily a pest of stored products, that feeds inside the grain as larvae inducing significant economic loss in various stored commodities. Our previous studies proved that garlic essential oil and its active substances inhibit oviposition in moths. To further explore the effect on reproductive behavior and accurately interpret the results in terms of effective control of the moth population, moths were treated with diallyl trisulfide (DATS), an active substance of garlic essential oil, at a dose of 0.015 µL/L in air (LC20, sub-lethal concentration). The results showed that fecundity and the proportion of viable eggs significantly decreased when the moths were treated with LC20 DATS. Furthermore, female circadian mating rhythms and calling periodicity changed significantly after treatment. Compared with controls, the peak in mating occurred approximately 1 h earlier on the first day after DATS treatment, while it was significantly later on days 2 and 3. Moreover, mating frequency declined in presence of DATS compared with the controls. The percentage of females engaging in calling behavior decreased significantly with time, to less than 50%, 2 days after treatment, while a high calling percentage (>80%) was recorded for control moths on all 4 days. In addition, DATS had an inhibitory effect on the mating duration of S. cerealella. Moreover, a significant reduction was observed in the amount of sex pheromones extracted 8 h and 9 h after treatment. Our findings suggested that DATS has the potential to manipulate the moth population at LC20 and would be an efficient alternative to synthetic insecticides for the control of pests having low toxicity to non-target organisms and ecosystems.

Do Helicoverpa armigera Moths Signal Their Fecundity by Emission of an Antagonist?

Female Helicoverpa armigera emit a pheromone, comprised of a 98:2 ratio of (Z)-11-hexadecenal to (Z)-9-hexadecenal, to attract males. It has been proposed that "immature" female H. armigera modulate attraction of males by emitting an antagonist, (Z)-11-hexadecenol, along with pheromone during the first two nights of calling. However, it is unclear why females would call and simultaneously release pheromone and an antagonist. We conducted observations of female calling during the first five nights after adult emergence to determine periodicity. We also measured the relative abundance of (Z)-11-hexadecenol to the major component, (Z)-11-hexadecenal, on the surface of the gland of calling females and compared it to the ratio of these two compounds inside the gland over the first three nights after adult emergence to determine how much antagonist may be released. We found that young females (< 1-d-old) are unlikely to call and, based on the relative proportion of (Z)-11-hexadecenol on the gland surface, even if they did call would be unlikely to release sufficient (Z)-11-hexadecenol to diminish male attraction.

Stink Bug Communication with Multimodal Signals Transmitted through Air and Substrate

This review represents complex mechanisms and processes of multimodal communication in stink bugs. During reproductive behavior the airborne and substrate-borne signals enable mate recognition, mediate directionality of movement, eliminate rivals and motivate partners for copulation. Species specific characteristics prevent hybridization at various levels of mating behavior. Male sex and/or aggregation pheromones as uni- or multicomponent signals attract mates to land on the same plant and there, trigger females to call males by vibratory signals, transmitted through the plant. Communication during courtship runs at short distance with visual, airborne, substrate-borne and contact chemical and mechanical signals. Abdomen vibrations produce the main repertoire of female and male calling, courtship and rival vibratory signals. To increase their informational value, stink bugs tune signal frequency, amplitude and temporal characteristics with mechanical properties of plants. The airborne component of species non-specific and high amplitude signals, produced by body tremulation and wing buzzing enables communication contact between mates standing on mechanically isolated plants. Female vibratory signals increase the amount of male emitted pheromone and the latter keeps female calling. Interaction, synergy and characteristics of visual, contact chemical and vibratory signals, exchanged during courtship remain under-investigated. Female and male competition for access to copulation in imbalanced sex conditions is characterized by duetting with rival song vibratory signals. Different receptors in and on different parts of the body are able to detect with high sensitivity multimodal airborne and substrate-borne communication signals. The relevance of the multimodal communication for the reproductive success of stink bugs is discussed.

Playback of male song advanced onset of nest building but not initial egg date or breeding synchrony in the Great Tit Parus major

Social stimulation can hasten and synchronize breeding in colonial species. However, whether it affects reproductive timing in territorial species, or whether social stimuli from different sexes are important remains largely unknown. Vocalizations of both sexes were played back in Great Tits. Females exposed to male songs built nests earlier, but did not lay earlier or breed more synchronously, while female calls had no effect. This study demonstrated social stimuli from males influenced nest building. In contrast to colonial species, laying date and breeding synchrony may be affected by other external cues in the Great Tits.