MALE HYPERSEXUALITY IN ECLECTUS RORATUS (ECLECTUS PARROT)
Tailai O’Brien, B.BUS. (HRM & Bus MGT), Cert IV Workplace Trainer, Cert III Captive Animals
Rob Marshall, B.V.Sc., M.A.C.V.Sc (Avian health)
Carlingford Animal Hospital, Sydney, Australia
Male hypersexuality in eclectus parrots refers to an ongoing state of unregulated sexual behaviour. Three presentations of male hypersexuality are described. The predisposition to this problem is the bird to human bond formed during the hand rearing process. Mating behaviours that characterise male hypersexuality include courtship displays, regurgitation, masturbation and aggression. These hormone-driven behaviours are stimulated by human interactions when the human bonded male bird reaches sexual maturity. The biologic basis of male hypersexual behaviour is the absence of a suitable breeding partner to bring an end to the mating phase of reproduction. The key features of the breeding biology of wild eclectus parrots that relate to this abnormal breeding behaviour are defined. Interactions and conditions in the home setting that stimulate male eclectus parrots to hypersexuality are described. Typical symptoms, the various forms of the condition, clinical implications, immediate treatment and long-term prevention plans are discussed.
Key words: Eclectus roratus, eclectus parrot, hypersexuality, unregulated sexual behaviour, breeding behaviour, cooperative, opportunistic, regurgitation, masturbation, aggression, reproduction, GnRH, GnIH.
Male hypersexuality in eclectus parrots refers to an ongoing state of unregulated sexual behaviour. This problem is managed by understanding the events that trigger the behaviour in the context of the biology in which they occur.1 There has been extensive scientific research into the reproductive biology of the Australian eclectus parrot (E.r.macgillivrayi).2 The results of these studies and observations in the aviculture environment provide the information needed to interpret and understand the abnormal social and sexual behaviours that typify this condition in the pet bird setting. Information regarding the unique breeding biology of eclectus parrots is provided in this paper as a means to understand the mechanisms of male hypersexual behaviour. We first describe the inbuilt reproduction systems that predispose hand-reared eclectus parrots to the unregulated sexual behaviours of male hypersexuality. Typical symptoms of this condition and its various forms are then discussed. Finally, clinical implications and treatment recommendations are noted.
The major biological influences on male hypersexuality in eclectus parrots relate to their distinctive breeding behaviours. These include cooperative breeding behaviour; seasonally opportunistic breeding behaviour and short highly regulated mating phase of the breeding cycle.
Cooperative breeding behaviour
The cooperative breeding behaviour of eclectus parrots is unique amongst psittacines.2 In the wild, nesting females receive all of their food (mostly fruit) from between one and five attending males who regurgitate it to them as pulp at the entrance of the nest tree hollow or on a nearby branch.2 Nestlings also get all of their food from the males, but indirectly via their mothers.2 In the context of male hypersexuality, this allofeeding behaviour is distinct from courtship feeding. Both terms refer to regurgitation of food by one individual to another. Courtship feeding is gonadotropin-releasing hormone regulated and testosterone driven breeding behaviour essential to successful insemination and male fertility.4 Courtship feeding and all other mating behaviours cease when the female lays eggs.5 After this time, regurgitation of food by the male is restricted to allofeeding behaviour. In this discussion, allofeeding refers to the nutritional support breeding behaviour that starts when the female occupies the nest then continues throughout the mating and parental care periods of reproduction and ends only when the female leaves the nest hole. In the wild, this feeding responsibility is often shared by several males and may continue for 8-11 months of the year.2 In the home setting, this same hardwired feeding behaviour is an accompanying feature of male hypersexuality. There are health consequences associated with male hypersexuality behaviours due to the substantial energy and immune cost of sustaining both courtship and allofeeding regurgitation over protracted periods of time.
Seasonally opportunistic breeders Eclectus parrots live in the tropics. They are seasonal breeders that adopt an opportunistic strategy to reproduction. This means they have distinct breeding and non-breeding periods but these are not necessarily consistent from year to year. Seasonal breeding activity in birds is regulated by the hypothalamic release of neuro-hormones particularly gonadotropin-releasing hormone (GnRH1) and gonadotropin-inhibitory hormone (GnRH1) to the pituitary gland in response to environmental cues.5 Key breeding prompts are integrated at multiple levels by GnRH1-receptive neurons with the effect of fine tuning the timing of reproduction. There is no synthesis of GnRH1 during the non-breeding period.5 Seasonal breeding birds detect environmental and climatic signals such as light (photoperiod), temperature, rainfall, food abundance and the presence of potential mates to time reproduction. These factors initiate the synthesis but not necessarily the release of GnRH1.5 Unlike temperate bird species, photoperiod and temperature have less influence on the breeding behaviour of tropical birds because temperatures and day-length across the tropics are relatively constant throughout the year. Food supply and climatic conditions have most impact on the time of reproduction for tropical birds. Conditions for breeding in tropical Australia regarding climate and food supply are ideal towards the end of the wet season. Most Australian tropical parrots breed in Autumn. For example, golden shouldered parrots (Psethotus chrysopterygius) time the start of reproduction according to the seasonal availability of green seed.6 The rainforest fruits that eclectus parrots eat are also in abundance towards the end of the wet season. Although conditions are considered ideal at this time, eclectus parrots delay the start of breeding until their nest hollows are suitably dry, which is usually in early spring. The start to breeding may vary considerably from year to year with the female occupying the nest any time between June and March. 2 In this respect, eclectus parrots can be classed as seasonally opportunistic breeders. Under the ideal conditions of a home environment, human-bonded male eclectus parrots will also take every opportunity to breed. Accordingly, male hypersexuality may occur for lengthy periods at any time of the year.
Rapid onset and short duration of mating phase of reproduction
The occupation of the nest hole by the female eclectus parrot is the ultimate stimulus that triggers the full functionality of the hypothalamic- pituitary- gonadal reproductive axis.
Breeding behaviour is initiated by the hypothalamic release of GnRH1.5 Until this time eclectus parrots in the aviculture environment show no signs of breeding behaviour. It is not known if GnRH1 is synthesised before this time. In response to GnRH1, the pituitary gland synthesises and secretes the gonadotropins luteinizing hormone (LH) and follicle stimulating hormone (FSH) into the bloodstream. LH travels to the gonads, where it stimulates the production of reproductive steroids such as testosterone and estradiol, whereas FSH guides gamete production. Given their importance in reproduction, it is not surprising that courtship and mating behaviours in males and females are modulated by gonadal hormones.5 The onset of breeding behaviour in eclectus parrots is rapid. Mating can occur within 24-48 hours of the female occupying the nest hollow. The female indicates mating receptivity by “begging” the male for food. The male responds by feeding her food, which marks the beginning of the mating stage of reproduction. This includes courtship displays and repeated allofeeding. Courtship behaviours culminate in copulation. Eclectus parrots mate during the day between the morning and afternoon meals. Copulation in captivity tends to be a lengthy affair compared to other birds. Eclectus parrots may copulate for up to 6 minutes before the male is satisfied by ejaculation, which can be heard as a faint whistle. In the wild, copulation appears to be a much shorter process. In established captive breeding pairs, the first egg of two may be laid as early as 7 days after the first mating.
Courtship rituals and copulation are regulated by GnRH1 and driven by the steroid hormone testosterone.7 Testosterone not only plays an important role in gamete production but also influences social and aggressive behaviour.7 Aggressive interactions often vary seasonally. They are typically high during the mating phase and decrease with the onset of parental care behaviour.7 Elevated testosterone is not maintained for prolonged periods because it can be costly leading to immune suppression and even increased mortality.7 Therefore, testosterone secretion is thought to be tightly regulated during breeding. The sex steroids provide feedback to the brain and pituitary, creating a regulatory feedback system necessary for reproduction and associated behaviours.5 In birds, neural inhibition of gonadotropins was thought to be solely the result of increased feedback into the brain from the pituitary and gonads. The discovery of GnIH and its active inhibitory effects of the reproductive axis has altered this view.5 GnIH is most active in males and females on the first day of incubation and first day after the first chick hatches.5 GnIH activity inhibits testosterone secretion either by directly acting on the gonads or acting within the brain.5 Female eclectus parrots reject the male’s sexual advances immediately after the second egg is laid. This response is thought to stimulate the release of GnIH, which lowers testosterone levels and rapidly extinguishes the sex drive of the male.5 The reaction facilitates the transition from aggressive and sexual behaviours to those relating to parental care. The immediate cause of male hypersexual behaviours in eclectus parrots is the absence of a suitable response from a female breeding partner to bring an end to the mating phase of reproduction. Without GnIH regulation, testosterone levels remain elevated resulting in uncontrolled mating behaviours and aggression. Male hypersexuality is the behaviour consequence of an unregulated and prolonged mating stage of the eclectus parrot breeding cycle.
A pivotal cause of this common condition is the pet bird to human bond formed during hand rearing. Upon reaching sexual maturity, hand reared males spontaneously seek out a potential mate. In the absence of a strong bond with members of their own kind they will pursue members of the human family as potential mates. They perform courtship advances that follow the same physical displays as wild birds. At first there is close face to face visual contact, head and body swaying, iris constriction and rumbling vocalisations. These preliminary courtship displays often go unnoticed by the human family. A perceived pair bond is formed when the preferred human(s) unknowingly responds to these advances in a physically affectionate manner. Kissing, hugging, cuddling and other forms of touch are common inadvertent reciprocal responses of human family members to the intense sexual interest being displayed by the male. The pet male recognises this positive human response as confirmation of his acceptance as a mate. In response, mating rituals are performed in the same way as wild males would compete for the female at the nest. Male courtship behaviour in eclectus parrots is initiated by the sexual receptivity of the female. As previously discussed, GnRH1 initiates mating impulses and GnIH ends them. GnRH modulation then regulates the sexually driven mating behaviours of the male.5 The mating phase of reproduction in eclectus parrots is sudden in onset and of short duration. With favourable seasonal conditions in an aviculture environment, mating behaviour occurs within 24-48 hours of opening the nest hollow. Eggs are laid 5-7 days later. GnIH brings the mating phase of reproduction to an end when the female starts to incubate her eggs.5 The mating period is of short duration often reduced to 10 days under aviculture conditions. Courtship and copulation behaviours are often ongoing in the hypersexual context because there is no such regulation of the mating period for human pair bonded males in the home setting. We consider the neurohormonal basis of male hypersexuality in eclectus parrots is unchecked elevation of testosterone levels caused by a failure of GnIH release.
Male hypersexuality in eclectus parrots is characterised by regurgitation as courtship feeding and/or allofeeding; copulation in the form of masturbation and aggression as a result of elevated testosterone or non-sexual territorial defence. Regurgitation and/or masturbation behaviours are directed towards human or inanimate surrogate mates. Inanimate surrogate mates may include corn cobs and similar sized solid food pieces, cushions, TV remote control devices, hanging plastic toys and any other firm object large enough to manipulate under the male’s body. A failure of the surrogate human or inanimate mate to reject the male’s sexual advances is responsible for the uncontrolled breeding behaviours that typify male hypersexuality namely courtship and allofeeding regurgitation, masturbation and aggression because there is no GnIH message to bring an end to the mating phase of reproduction. Without this neurohormonal message, the male remains in an unrestrained testosterone driven mating state.
We refer to three clinical presentations each reflecting their own variation of natural breeding behaviours. The first incorporates the full sequence of mating behaviours; the second is restricted to masturbation; the third involves regurgitation in the form of allofeeding.
The first presentation occurs in males that have yet to establish a permanent pair bond with a single member of the family or inanimate surrogate mate. This common form of male hypersexuality includes the full assemblage of mating behaviours expressed by wild males, namely courtship displays, courtship feeding followed by copulation in the form of masturbation. Allofeeding also takes place. This presentation is the most troublesome in regards to health and wellbeing because the complete array of male breeding behaviours is expressed including allofeeding regurgitation. The second presentation involves masturbation alone. This behaviour is usually seen in mature males (older than 6 years) that have established a strong and unbroken pair bond with a particular human family member over a number of years. The third form of male hypersexuality presents as allofeeding regurgitation without any other breeding behaviours. This uncommon condition is usually confined to homes with more than one male bird. In this situation, alpha birds subordinate lesser males to feeding support status as happens in the wild.9 These lesser birds exhibit non-sexual allofeeding regurgitation rather than the testosterone driven courtship regurgitation behaviour of the household’s alpha male.9 Although not strictly sexual behaviour, allofeeding is included in this discussion as a breeding behaviour commonly involved with male hypersexuality.
Aggression is a prominent feature of all presentations. For the first two presentations, aggression is driven by elevated testosterone. In regards to allofeeding, aggression is territorial rather than a sexual response. Aggressive encounters (challenges) can also result in an increase of circulating testosterone, which in turn facilitates aggressive responsiveness.8 Extreme aggression in households where several family members are competing for the attention of a human surrogate mate reflects this type of biological reaction. Testosterone may be immunosuppressive but only in males in poor condition that cannot afford to invest in reproduction as well as immune function.8 These findings suggest that males with the highest quality sexual signals during the breeding season, at least partly due to high testosterone levels, are advertising their superior immunocompetence to females.8 The immunosuppressive consequences of elevated testosterone are commonly encountered with male hypersexuality probably because captive eclectus parrots are generally genetically and physiologically inferior to their wild conspecifics.
In the clinical setting, male eclectus parrots may be presented for the behavioural consequences of male hypersexuality, namely regurgitation, masturbation and aggression, or health problems associated with elevated testosterone or the impact of stress from these behaviours. Regurgitation and masturbation are rarely the presenting complaint. Aggression in previously calm affectionate males is of concern to most owners but the large majority of birds are presented with hypersexual-related health problems. Owners rarely recognise the connection between the health episode and hypersexual behaviour. In eclectus parrots, health issues associated with this condition include digestive disturbances, and feather destruction associated with xerosis, dry feathers due to decreased bathing frequency and aged feathers as a result of delayed moult. The pathogenesis and description of these conditions is outside the scope of this paper. Once established it is extremely difficult to cure the sexual regurgitation and masturbation behaviours associated with male hypersexuality without interfering significantly with the emotional relationship the owner experiences with their pet bird. Our methods for managing male hypersexuality focus on prevention and early intervention. Establishing a daily routine from a young age combined with continuous behavioural training and sufficient exercise is critical to support a reciprocally rewarding bond focussed on having fun throughout all the stages of life. Learning, playing, mutual affection and respect are critical elements of positive behavioural patterning that eliminate the development of bird to human interactions which are characterised by sex behaviours in adult companion parrots.
Immediate treatments are listed below. These recommendations are combined with treatments targeting health issues associated with male hypersexuality.
Withdraw physical contact temporarily for 48 hours. After this time do not allow the pet bird to reciprocate touch. This is achieved by redirecting attention away from mutual grooming and towards independent activities such as foraging.
Pre-empt and eliminate all regurgitation and masturbation attempts on the human, or inanimate objects permanently. Send a strong message to the male verbally with a harsh command. Return the bird to its cage immediately when regurgitation is attempted.
Remove all familiar items and toys in the cage that are used for the purpose of regurgitation or masturbation. These items must be replaced with novel toys that engage the brain. Redirect the bird’s attention away from the sexual bond with their owner. This is best achieved by establishing a daily routine centred around morning and afternoon mealtimes to encourage natural biorhythms and natural bathing behaviour. Daily showers or mist sprays will promote appropriate, time consuming grooming activity. A drenching bath will expend most of the bird’s pent-up energy.
Meaningful and interactive occupation activities, involving learning activities and training will help divert the bird’s attention from sexual gratification. Encourage daily exercise and supervised flight around the home. Daily training must be scheduled for side by side interaction and rewards for appropriate behaviour.
Van Sant F. Problem sexual behaviours of companion parrots. Manual of Parrot Behaviour ed. AU Luescher. 2006; 233-244.
Heinsohn R. Ecology and evolution of the enigmatic eclectus parrot, Eclectus roratus. Journal of Avian Medicine and Surgery. 2008; 22:146-150
Fusani L. Testosterone control of male courtship in birds. Hormones and behaviour. 2008; 54:227-233
Spoon TR. Parrot reproductive behaviour. Manual of parrot behaviour. 2006; 8:63-74
Ubuka T. Neuroendocrine regulation of gonadotropin secretion in seasonally breeding birds. Frontier of Neuroscience. 2013; 38:323-348
Garnett S. Golden-shouldered Parrot. Nature Australia. 1999; 25, 18-19
Swett MB. Interaction of testosterone, corticosterone and corticosterone binding globulin in the white-throated sparrow. Comp Biochem Physiol. 2008; 151:226-231
Roberts M. Is testosterone immunosupressive in a condition-dependent manner? An experimental test in blue tits. Journal of Experimental Biology 2009; 212:1811-1818
Pikus A. Testosterone social status and parental care in a cooperatively breeding bird. Hormones and behaviour. 2018; 97:85-93
Marshall R. Exploring the wild diet to solve functional digestive disorders in eclectus parrots. ICARE. 2017; 497-506
Stomach Dysfunction in Eclectus Parrot - ICARE Conference April 2019
Audio visual presentations from 2018 Atlanta AAV Conference