European Food Safety Authority — Safety and efficacy of Panaferd-AX (red carotenoid-rich bacterium Paracoccus carotinifaciens) as feed additive for salmon and trout

Safety and efficacy of Panaferd-AX (red carotenoid-rich bacterium
Paracoccus carotinifaciens) as feed additive for salmon and trout1
Scientific Opinion of the Panel on Additives and Products or Substances used in Animal Feed

(Question No EFSA-Q-2006-173)
Adopted on 18 September 2007

PANEL MEMBERS
Georges Bories, Paul Brantom, Joaquim Brufau de Barberà, Andrew Chesson, Pier Sandro Cocconcelli, Bogdan Debski, Noël Dierick, Anders Franklin, Jürgen Gropp, Ingrid Halle, Christer Hogstrand, Joop de Knecht, Lubomir Leng, Anne-Katrine Lundebye Haldorsen, Alberto Mantovani, Miklós Mézes, Carlo Nebbia, Walter Rambeck, Guido Rychen, Atte von Wright and Pieter Wester

SUMMARY
Following a request from the European Commission, the Panel on Additives and Products or Substances used in Animal Feed (FEEDAP) was asked to deliver a scientific opinion on the safety and efficacy of Panaferd-AX. The product is a feed additive consisting of dried sterilised cells of a red carotenoid-rich bacterium (Paracoccus carotinifaciens) intended to provide farmed Atlantic salmon (Salmo salar) and rainbow trout (Oncorhynchus mykiss) with a source
of astaxanthin which confers the characteristic red colour to the flesh. Panaferd-AX contains around 4 % total red carotenoids, predominantly astaxanthin (2.2 %), adonirubin (1.3 %) and
canthaxanthin (0.4 %).

In salmon and rainbow trout astaxanthin, deposition in flesh from Panaferd-AX was less efficient than that from synthetic astaxanthin. However, equal astaxanthin doses from both sources resulted in a comparable flesh pigmentation in the dose range of 20 to 100 mg astaxanthin kg-1 feed, due to the contribution of the other red carotenoids, mainly adonirubin and canthaxanthin, which are also demonstrated to be deposited in fish flesh. The technological
and organoleptic properties of flesh from Panaferd-AX-treated fish were not different from those treated with synthetic astaxanthin.
Panaferd-AX, at dietary incorporation rate of 12.5-fold greater than the proposed maximum incorporation rate (0.4 %) is safe for salmonids (trout and salmon). Paracoccus carotinifaciens is not a known pathogen, and no other concerns have been identified either in the limited literature available or in the data submitted in the dossier.
. . .
Click HERE to download the report as a PDF.
http://www.efsa.europa.eu/sites/default/files/scientific_output/files/main_documents/feedap_op_ej546_panaferdax_slm_tt_en,3.pdf

Marc Weiss’s pair of Swainson’s toucans

Swainson’s toucans

Marc Weiss bred this pair of Swainson’s toucans.

As a protein supplement, Marc fed the birds hard boiled chicken egg not mixed with bread crumbs. The toucans used a wooden box for a nest. A palm log also was in the cage. The birds demolished this and used some of the debris in the box for nesting material. Marc Weiss believes that the “trashing” of the palm trunk is an important part of nesting behavior.

Here’s what Marc Weiss wrote about the birds care:
Their basic diet was fruit, and Mazuri softbill diet. Breeding was induced using my egg food, not just egg. Mixture of mashed eggs, soy protein isolate and vitamin/mineral mix. They would also eat anything living that got into the aviaries which had half inch X two wire as I remember. So any bugs, lizards, etc. would be eaten as they were outdoors.

Swainson’s toucan chick

POTENTIAL FOR STIMULATING REPRODUCTION IN CAPTIVE BIRDS BY PHOTOPERIOD MANIPULATION

By Arthur C. Risser, Jr., Ph.D.

IFCB 1978

There is no more regularly occurring event in nature than the changing of daylight to darkness. Since the beginning of time, this event has had a profound influence on the seasonality and cyclicity of all organisms. Birds in general and north temperate birds in particular have evolved to gear their annual activities to seasonal changes and specifically to those changes related to day length. Seasonal day length changes provide the most reliable information indicating the approach of a suitable breeding season. However, photoperiodic responses are modified by many environmental inhibitors and accelerators such as temperature, weather conditions, food supply and behavioral interactions. The photo refractory phase of the annual breeding cycle which is present in many, but not all species, acts as a safety mechanism to prevent unseasonal breeding.

Ever since Rowan’s 1920 – 1930’s early work with Juncos in Alberta, Canada, scientists have been intrigued with the physiological mechanisms and the behavioral correlates to changes in day length. Rowan’s early work demonstrated beyond a doubt that increasing the number of daylight hours after the short photoperiod of fall and winter was stimulatory to the migratory and reproductive responses. With added illumination, he was able to trick his captive Oregon Juncos into breeding in the dead of winter even when there was snow on the ground. It is now standard procedure for poultry people to hasten sexual maturity and maintain production by manipulating the lighting scheme in the hen house. Those of us dealing with captive stock should, where possible, consider utilizing this powerful stimulus as a tool to promote reproduction in our captive birds.

This report will give a very brief and general summary of the vast literature which exists relative to photoperiod, discuss some situations in which this type of manipulation has worked or is work= ing and then mention how we as aviculturists may be able to utilize light manipulation to our advantage. This summary is based on two excellent reviews. (See Lofts and Murton, 1968, Zoological Society, London, 155, 327-394; Farner and Follett, 1966, Journal Animal Science, 25, supplement.)

We could be very teleological and state that a bird’s primary purpose in nature is to maximize the offspring it leaves. To do this, a species must accumulate enough energy to make eggs and then provide food to the young that hatch. To achieve breeding condition and anticipate a season of optimum food availability, birds have evolved response mechanisms to various appropriate environmental stimuli which signal the approach of a suitable season. It is, therefore, recognized that the increasing day length which is stimulatory to a great many species is tempered by other factors such as temperature and rainfall which can lead to the emergence of an important dietary item. While increasing day length is stimulatory to gonadal development, it is equally true that under the influence of long summer days, gonads of many species spontaneously regress as the hormone secretions from the anterior pituitary gland diminish and bring about a period of sexual quiescence called the “refractory period”. It has also been demonstrated, however, that some species of birds lack a true refractory condition.

Evidence for gonad stimulation by light and the following refractory period has come through the efforts of physiologists and field ecologists working first with natural populations and then with captive birds to characterize the gonadal cycles of a variety of species. These investigations have meant either harvesting a representative sample of birds from a population at a given latitude, measuring the gonads (or later, examining the bird’s gonads by laparotomy and putting them back into the population). Then light manipulation experiments (especially on White-crowned Sparrows, House Sparrows, starlings and House Finches) helped elucidate the photoperiod mechanism in operation. Some species received more attention than others because that particular species was superabundant or an economic pest, and investigations into breeding cycle might provide information on how to shut off its breeding. Probably sixty to seventy north temperate species in widely separate orders have been shown experimentally to depend on light stimulation to bring about gonadal recrudescence. But where the male may make a full response to increases in day length and undergo active spermatogenesis, females may not give full response and may depend on other factors (i.e., stimulus provided by presence of mate, nesting site, etc.) for full breeding expression.

Basically, the mechanism bringing about the gonadal response is as follows. Appropriate day length is perceived by external receptors and transmitted to the hypothalamus of the brain where the information is further processed. A circadian or 24-hour clock – which is still largely a mystery – influences the hypothalamus, and special chemical signals are then sent to the anterior pituitary which, in turn, releases gonadotropic hormones (FSH and LH) which signal the gonads to undergo egg and sperm production. In turn, the gonads produce sex hormones which bring about certain secondary sexual characteristics and the appropriate reproductive behavior.

One interesting aspect coming out of experimental work is that(at least in some species) the photoperiod response depends on stimulation being received during a light sensitive stage of the circadian rhythm. In other words, light stimulus provided at the wrong time in the species’ 24-hour rhythm may have no response.

There is also considerable field evidence gathered by ecologists to indicate that the breeding cycle may be modified by such environmental factors as temperature, weather and food, but none of these factors bring about any gonad response in the absence of proper light stimulation. Temperature (and even more importantly, a mild winter) has pushed several European species through a winter breeding season when normally they would be inhibited because of poor weather conditions. It is thus important to note that not only is increasing day length stimulatory but so also is the decreasing daylight of late fall and winter to which some species would respond minimally. Under extremely favorable conditions it is conceivable that this autumnal response could become productive. Usually, however, a variety of environmental factors are inhibitory and shut down this autumnal surge before it is completed.

Complicated behavioral factors may also influence the final stages of breeding. Apart from requiring the right habitat and the possession of a territory, most birds depend on a host of reciprocal displays between mates to synchronize and regulate the final stages of reproduction. Males may initiate the breeding season by assuming a reproductive condition before females. The final timing of the season (oviposition) depends on the female receiving the appropriate stimuli from the male and her environment. Stereotyped behaviors can cause specific hormone secretions, a fact well worked out by several studies conducted mainly on captive doves and canaries, although there are still countless details to be settled.

But what of the refractory period or period of sexual quiescence? Anatomically, this period is marked by rapid degeneration of gonadal activity and the accumulation of cholesterol-rich fat tissue in the testes. It involves a condition during which hormone response cannot be brought about by photo-stimulation and has probably evolved to prevent breeding at inappropriate times. (In captivity, however, our objective is to try to provide optimum conditions on a year-round basis). Once a bird emerges from its refractory state, it is once again photo-sensitive. The duration of this refractory period varies among species. It can be as short as six weeks (Quelea) or as long as three or four months. Many temperate zone species, i.e., the mallard, regain their photo-sensitivity in the autumn but remain sexually inactive due to such inhibitory factors as cold temperatures, lack of food and diminishing day lengths which are prevalent in their environment.

In the bobolinks (a trans-equatorial migrant) it has been shown that exposure of autumn birds to an eight-week period of ten hours daylight before stimulating with fourteen hours of light will advance the development of breeding condition by two months over birds maintained on fourteen hours daylight all the time. Trans-equatorial migrants encounter refractory-breaking day lengths in crossing the equator, but since it takes time to go through a refractory phase, southern hemisphere breeding by such birds is an exception rather than a rule. Moreover, some species may lack a photo refractory period entirely and thus be capable of being artificially stimulated throughout the year (as with some species of quail).

It is, therefore, important to emphasize that in any photo-periodic manipulations on captive birds we must be cognizant of this refractory period and use shortened periods of exposure to light as a means to dispel it.

Experiments by Wolfson (see Photoperiodism and Related Phenomenon in Plants and Animals, 1959, AAAS.) have shown that the duration of the daily photoperiod determines (1) the rate at which testicular growth and spermatogenesis proceeds and thus the time of year when it occurs, (2) the degree of response and (3) the duration with which a response persists before the refractory phase begins. Furthermore, the refractory period is dependent on the photoperiod in a quantitative way, short photoperiods hastening its early dispersal and long ones prolonging it.

The importance of photoperiod varies from place to place. Tropical and equatorial breeding cycles are considerably different from those of temperate species. In high latitudes (northern North America, Europe and the Arctic Circle) photoperiod is the main regulator of sexual cycles in many species but in low latitudes, where seasonal changes in day lengths are less, the position is not quite so clear, and factors in the immediate environment are often more important. According to investigations by Alden Miller (formerly Museum of Vertebrate Zoology, Berkeley) in equatorial latitudes (5° south – 5° north), photoperiodic fluctuations may have little or no influence on birds’ reproductive rhythms, but between 5° and 10° north and south latitude, a prevailing influence of photoperiod can be detected.

In equatorial and tropical environments, where annual fluctuations in daily photoperiod are slight or absent, seasonal rains and associated changes in vegetation act as the environmental synchronizers to reproduction. In Indonesia, for example, most species, including evergreen inhabitants, breed from January to March in West Borneo and March to July in Java, depending on the rainy season in the two areas. In contrast, Central American birds mostly breed between April and June, before or early in the seasonal rains, while in southwest Ecuador breeding begins in the onset of rains in the December – April period. Many African land birds have breeding patterns associated with rainfall and show a variety of preferences in different areas. Several Galapagos Finch species brought back to California and held captive by Dr. Robert Orr at the California Academy of Science were stimulated to sing when artificial rain fell on their cages. It is highly likely that the tropical situation reflects a relative insensitivity to and perhaps lack of dependence on photo stimulation. However, some species may still have the capacity to respond to photo stimulation as well as to other types of stimulus. The trick is to determine what the proper stimulation is. A similar breeding response to sporadic rainfall has been noted in many arid or semi-arid species in Australia so that approximately thirty out of sixty species examined in detail attained full breeding condition within a month of rainfall having occurred. Moreover, Australian semi-arid birds studied experimentally showed that they can be affected by variable photoperiods, though under natural conditions other stimuli seemed to be stronger and capable of overriding the effects of light. These opportunistic species have also evolved the ability to respond quickly to suitable breeding conditions by undergoing rapid gonadal maturation as juveniles.

Although not always the case, domestication and captive conditions can have a definite influence on how responsive species will be to photoperiod changes. The common pigeon has had a domestic history dating back perhaps to 3,000 B.C. Under the proper stimulatory conditions, common pigeons (and some of their well-known relatives such as the Collared Dove) may have no refractory period. This may also be true of the Red Jungle Fowl and certainly of some of its phylogenetically close relatives (e.g., some quail).

But what of captive conditions where we have been known to move birds from the southern to the northern hemisphere and vice versa. In general, those species with restricted photoperiodically controlled breeding seasons modify their cycles to suit the day length changes in their new environment.

It is often the case that for the first season an unseasonal response is shown, presumably because the hormone apparatus has been stimulated inappropriately during transfer to the new environment. Thus, we may find southern hemisphere birds coming out of quarantine and laying eggs immediately upon entry into a new aviary condition. (Then there is the case of Silver Gulls taken from Australia to a zoo in the United States. For two years the birds nested in November, then changed to breeding in spring and summer. Seemingly more remarkable is that after twenty years of captivity their descendants reverted to autumn breeding.) Other species such as the emu, budgerigar and European Stork when moved from one hemisphere to another, retain their original breeding season despite a change in the light regime. Other species when moved across the equator, however, shift their breeding season by six months, suggesting that their inherent annual reproductive rhythm can indeed be modified.

Given the condition under which many zoos and private aviculturists keep captive birds, day length is simply one variable which can be controlled. With indoor bird exhibits or housing situations where birds are kept indoors all the time or for a large portion of the year, manipulation of photoperiod can be an easy matter. Timing clocks to turn lights on and off at predetermined intervals are inexpensive and can even be purchased with mechanisms which will automatically change light period on an annual basis to correspond with any predetermined latitude. With today’s modern solid state electronics, it is further possible to provide lighting systems which mimic normal dawn and dusk twilight periods. Ability to provide dawn and dusk twilight is particularly important in arboreal species but not so important for ground dwelling species.

Several captive breeding programs are currently employing photo-period manipulation in their reproductive programs. (1) Patuxent Wildlife Research Center: Whooping Cranes and Masked Bobwhite, (2) Bronx Zoo: Puffins, (3) Cornell University: Peregrine Falcons, (4) Sea World, San Diego: Penguins.

While photoperiod manipulation offers an opportunity to enhance or modify reproduction in captive birds, the following suggestions are offered. (1) Become familiar with the natural history of the species to be subjected to light manipulation. Not all captive birds respond the same. Not only are there species differences but also individual differences within the species. (2) So nearly as possible, try to determine where the captive stock came from. Different populations may have evolved with different conditions. (3) Try to set up replicate experiments with controls. Repeatable experiments lend statistical validity to any results. This may be particularly hard since with some of our exotic birds the sample size is very small.

From an avicultural point of view, day length manipulation may have several advantages. (1) The captive environment can be made to mimic more exactly the conditions under which the species lived in the wild. It is quite possible, therefore, to influence the time at which breeding starts. (2) It may be possible to force a species into a seasonal reproduction and thereby stagger husbandry efforts. (3) By initiating breeding earlier, it may be possible to promote more than the usual number of clutches in a season.

BREEDING STRATEGIES IN ESTRILDID FINCHES OF DIFFERENT CLIMATE ZONES: EFFECTS OF ULTIMATE AND PROXIMATE FACTORS

Granted by the Deutsche Forschungsgemeinschaft

Dr. Roland Sossinka
Universitat Bielefeld
West Germany

IFCB 1978

INTRODUCTORY REVIEW ON BREEDING PERIODICITY IN BIRDS
Birds are dependent upon environmental factors for successful reproduction more than other classes of vertebrates. “All species of birds breed at those times of the year when, on the average, young can be profitably raised (Lack 1950). This statement sums up a great number of observations and investigations on breeding periodicity and related ecological factors in many species of birds. The general correctness of Lack’s rule raises two questions: (1) Why are the birds programmed to breed only at those times when young can be profitably raised? and (2) How does the bird “know” when to breed; that is, in what way is the timing of breeding periods accomplished?

The first question deals with the ultimate control of breeding periodicities and the second with the proximate control. With the exception of very few habitats (e.g., tropical islands, Ashmole 1968), in most places on earth there is a succession of periods more favorable to reproduction and periods when reproduction is impossible (or at least requires much more energy input with a decreased probability of success). The duration of the favorable conditions can be very different: three quarters of a year, as in many tropical areas, or one month only, as in some subarctic places. The favorable period can reappear at regular intervals, linked to annual seasons (for review, see Immelmann, 1971), or irregularly, as in some deserts with sporadic rainfalls (Serventy, 1971). The favorable period is characterized by the presence of that environmental factor or combination of factors which actually guarantee the breeding success. These factors are called “ultimate factors” (Thomson, 1950). In many populations of birds, food is the most important factor. After the young are hatched, much more food is needed than before and often a special type as many species raise their young with food different from their normal adult diet (Marshall, 1951). That is why species of the same area, specialized on different food sources, often differ in their breeding periods.

Further ultimate factors are the abundance of nesting sites and nest material, protection against predators, nd the absence of adverse climatic factors such as heavy rainfalls or low temperatures. During the phylogeny, each species had its breeding period restricted to those times of the year when these ultimate factors were present. Individuals who tried to reproduce outside the favorable period had much higher energy costs and were less successful. They gave their genes – which had information for not restricting reproduction to the favorable season – to their young which, born in the absence of the ultimate factors, had a reduced chance of survival and reproduction. That is why selection favored those genomes which restricted breeding to the favorable times of the year only.

While the ultimate control acts during evolution, the proximate control of breeding periodicity takes place in the individual. The birds, whose gonads are in a state of quiescence outside the breeding period (to prevent them from breeding at the “wrong” time and to conserve energy, e.g. by reducing weight during flight) have to activate the entire sexual system. This means that the hypophysial-hypothalamal axis has to release gonadotropic hormones which activate the gonads. The ovary and testes must increase in size, the germinative cells undergo maturation, and sexual hormones must be produced. The latter substances (in some species gonadotropins also) cause some changes in seasonal secondary sex characteristics (e.g., bill coloration or nuptial plumage) and are responsible for the appearance of a variety of sexual behavior patterns. All these events have to take place before the onset of the favorable season. There are two ways to start these preparatory processes in time: (1) External factors trigger the activation or (2) Internal factors (endogenous rhythms) are responsible for the activation.

If any external factor shall start the gonadal activation, this factor must be noticeable to the bird and must occur at a certain interval before the onset of favorable conditions. It has to predict highly reliably the appearance of the ultimate factors in advance.

If such an external factor exists, then during evolution each species’ respective population will develop response mechanisms to these predicting factors which then are called “proximate factors” (Thomson, 1950).

One of the most important proximate factors in birds living in the moderate climate zones is the photoperiod. Many species and subspecies of birds respond to a certain population-specific length of daylight with gonadal recrudescence. Twelve to fourteen hours after sunrise a short light sensitive phase occurs in each individual which, stimulated by long day conditions, is able to transform this stimuli (perceived by the eyes or by extraocular receptors in the central nervous system) into neuro-secretory activity of some specialized cells in the hypothalamus, which results in the release of gonadotropic hormones (Farner, 1970). In the more equatorial zones, the resident species of birds often have a reduced or no photosensitivity. Other environmental factors are known to act as proximate factors; for example, rainfall and related phenomena (Moreau, 1950).

If the favorable season reappears regularly, the gonadal activa- tion can be started by internal factors. An endogenous circ-annual rhythm can induce gonadal maturation at intervals of nearly one year. This was proven in birds kept in laboratories under absolutely constant conditions (Gwinner, 1969). Their endogenous cycles, running free in constant illumination, had an average period length of eight to eleven months. In feral birds this endogenous rhythm becomes synchronized to the annual seasons by external factors (proximate factors in a wider sense), which then are called “Zeitgeber” (Aschoff, 1954). In many of the species of birds investigated thus far, internal and external factors both take part in the regulation of breeding periodicity.

Each population has evolved a pattern of reproductive activity and inactivity and mechanisms regulating the onset and the end of gonadal activity which are highly adaptive, according to the climatic conditions of the area from which it is derived.

MATERIAL AND METHODS
To investigate some parameters of sexual maturation, we selected three species of Estrildid Finches from very different climatic zones.

The family of Estrildidae is known to be a well-defined group of closely related genera (Delacour 1943). Therefore, one should expect to find a generally uniform type of sexual maturation, probably modified by some species specific adaptations. The Estrildids originated in the tropical areas of mid-Africa. From there they dispersed in several waves to India, Southeast and East Asia and to the Australian region (Delacour, 1943, Mayr, 1968). The three species we investigated are: (1) the Fire Finch Lagonosticta senegala from Africa, (2) the Zebra Finch Poephila (Taeniopygia) guttata castanotis from Australia and (3) the Spice Finch Lonchura p. punctulata from India.

We restricted our investigations in the first study to the process of sexual maturation in young individuals, which had the advantage that the confounding factors of prior sexual activity and gonadal atrophy were not present. The ultimate control is the same in first year breeding birds as in adults, and most of the physiological mechanisms regulating the gonadal activation should be the same as well (Sossinka, 1970).
The birds were bred in indoor aviaries or cages. Non-domesticated or only slightly domesticated strains were used (Sossinka, 1970). The young were separated from their parents at about thirty-five days of age and were kept in bisexual groups under relatively constant laboratory conditions (light-dark cycle 14 : 10 hours, temperature 21 to 24° C. relative humidity 55 to 70%). They were fed a diet of mixed millets and seeds, sprouted millet and water ad lib, and three times a week a multivitamin emulsion was added to the water. Additional egg-food and insects (beetle-larva) were given during the first thirty-five days of life when nestlings were being fed by their par-ants.
At regular intervals body size, moult and gonadal size were checked. The following measurements were taken in young males: body lengths; wing length, bill length and width; body weight; number of renewed feathers; volume of the left testis, as measured by laparotomy with an ocular micrometer in a surgical microscope (Sossinka, 1974).

RESULTS
Body growth slows down in all three species after a continuous linear rise. Adult measurements are reached in the Spice Finch

(13 g body weight) and Zebra Finch (12 g body weight) about day 28 to 30 and in the smaller Fire Finch (8 g body weight) at about day 25. The onset of juvenile molt, when the first feathers of the adult plumage appear, is not later than day 35 in the Fire Finch and Zebra Finch and day 40 in the Spice Finch. The end of this juvenile molt, however, differs markedly between the three species. Here the principal differences in time patterns which are typical of the maturation of all primary and secondary sex characters can be seen. The Fire Finch finishes juvenile molt at about 150 days of age, the Zebra Finch around day 90, and in the Spice Finch it takes 200 to 300 days (in some birds even more than one year).

These three differing patterns – a medium, an accelerated and a retarded development – appear even more pronounced in the growth curves of testes volume. Plotted logarithmically, in all three species an initial volume increase in the nestlings is followed by a period of quiescence. Growth is resumed at different ages in each species, and the time needed to complete gonadal growth is very different also. Adult male testes volume is reached in the Fire Finch at 130 days, in the Zebra Finch at 70 days and in the Spice Finch at 250 days. The latter shows the largest intra-individual variation while the Zebra Finch shows the smallest (Figures 1 – 3). In Figure 1, the curve of testes volume in the Fire Finch is slightly corrected to equalize the differences in both weight. For absolute data, subtract 0.2 units.

DISCUSSION
In general, the type of gonadal maturation is the same in all three species, characterized by an intermediate phase of quiescence. This takes place at the same developmental age in the three species and is comparable to the juvenile refractory period in photosensitive species (Miller, 1954). Besides this homology, the species investigated showed pronounced differences in the rate of maturation. By considering the climatic conditions in the area of origin in each species, the selection pressures responsible for these adaptations become evident.

The Fire Finch lives in and at the edge f the tropical forest of Africa. As reported by Morel (1967), its breeding period is extended to nine or ten months a year with only a short restriction. Independent of the time of the year in which they were born, nearly all young birds can reproduce successfully themselves if their maturation is not too fast or too slow. Very fast maturation means high energy costs and the risk of inexperienced parents insufficiently foraging for food for the nestlings. Very slow means a disadvantage in competing with conspecifics. This presumably primitive pattern of a moderately rapid rate of maturation is not advantageous in central Australia where the Zebra Finch is found nor in India, the native habitat of the Spice Finch. Zebra Finches live in almost all parts of Australia, especially in the arid areas of western and central Australia. Here the rainfalls, which are essential for the food supply of the young, are rare and occur irregularly (Serventy, 1971). The Zebra Finch is an opportunistic breeder, starting to breed immediately after rain (Immelman, 1962). Only those young birds which have an extremely rapid rate of maturation can reproduce in the same vegetation period in which they were born. Otherwise, they have to wait until the next erratic rain falls, which in some places may not occur for twenty or more months. For this reason, natural selection has favored an extreme precocity in this species (Sossinka, 1970, 1974).

In the Spice Finch, on the other hand, egg laying in northern India takes place only during two months of the year (Thapliyal, 1968). Outside this strongly restricted season, successful reproduction seems to be nearly impossible. That is why young birds have to wait for three quarters of a year to start reproduction themselves and, therefore, their rate of maturation is retarded.

While in the Fire Finch and the Zebra Finch, under constant laboratory conditions, the rate of sexual maturation appears rather uniform, in the Spice Finch there are great intra-individual variations. This indicates the relatively environment-independent, internal type of maturation in the Fire and Zebra Finches in contrast to some possible influence of external factors in the Spice Finch. There is evidence that a cold and wet period acts as a proximate factor inducing the onset of sexual activation.

SUMMARY
The ultimate factors in the different climatic zones from which the species investigated come cause different patterns in breeding periodicity. According to these differences, the rate of maturation is quite different in the young of these related species: moderately
the in the tropical Fire Finch, extremely rapid in the precocious, opportunistic-breeding Zebra Finch, and retarded in the yearly breeding Spice Finch.

REFERENCES
Aschoff, J. 1954. Zeitgeber der tierisch Jahresperiodik.
Naturwissenschaften 41:49-56, 7
Ashmole, N.P. 1968. Breeding and molt in the White Tern (Gy is alba) on Christmas Island. Pacific Ocean, Condor 70:35-55.
Delacour, J.A. 1943. A revision of the subfamily Estrildinae of the family Ploceidae. Zoologica 28:69-86.
Farner, D.S. 1970. Day length as environmental information in the control of reproduction of birds. In: La photoregulation de la
reproductiion chez les oiseaux et les mammiferes. Ed.: Benoit
et Assenmacher, Paris, 71-91.
Gwinner, E. 1969. Untersuchungen zur Jahresperiodik von Laubsaingern. J. Orn i thol . 110:1-21.
Immelmann, K. 1962. Beitrage zu einer vergleichenden Biologie
australischer Prachtfinken. Zool. Jb. Syst. 90:11-196,
Immelmann, K. 1971. Ecological aspects of periodic reproduction. In:
Avian biology. Ed.: Farner, King, Parkes, New York-London.
Marshall, A.J. 1951. The refractory period of testis rhythm in birds and its possible bearing on breeding and migration. Wilson Bull. 63:238-261.
Mayr, E. 1968. The sequence of genera in the Estrilda (Aves). Breviora 287:1-14.
Moreau, R.E. 1950. The breeding seasons of African birds. I. Land birds. Ibis 92:223-267.
Morel, M.Y. 1967. Les oiseaux tropicaux elevent-ils autant de juenes qu’ils peuvent en nourrir? Le cas de Lagonosticta senegala. La Terre et la Vie 77-82.
Serventy, D.L. 1971. Biology of desert birds. In: Avian biology.
Ed.: Farner, King, Parkes. New York-London. I. 287-339.
Sossinka, R. 1970. Domestikationserscheinungen beim Zebrafinken Taeniopygia guttata castanotis (Gould). Zool. Jb. Syst. 97: 455-521.
Sossinka, R. 1975. Quantitative Untersuchungen zur sexuellen Reifung des Zebrafinken, Taeniopygia castanotis Gould. Verh. Dtsch. Zool. Ges. 344-347.
Thaplial, J.P. 1968. Body weight cycle of the spotted Munia, Lonchura punctulata. Bull. Nat. Inst. Sci. Ind. 86:151-166.
Thomson, A.L. 1950. Factors determining the breeding seasons of birds: an introductory review. This 92:173-184.

THE BREEDING OF PSITTACINES

By R. J. Topper

IFCB 1978

There is no such thing as one correct way in the handling, care or breeding of psittacine birds.

One of the key factors in the breeding of psittacine birds is nutrition. If we look into the investigations that have been done on these birds in their wild or native habitats, we will find that they are predominantly grain eaters. This is to the exclusion, for the most part, of seeds. We normally think in terms of seeds representing flowering type plants and grain representing non-flowering plants, which would include what we call grass seeds. However, we have to be cognizant of the fact that there is no such thing as a grain or combination of grains or seeds that can give a bird the total nutrient levels that it requires and has been able to obtain in its natural habitat. Certainly, birds eat many things; berries, fruits, nuts, grains, some seeds, grubs, etc. Realizing that we cannot duplicate these exactly, we certainly hope to be able to nutritionally or chemically duplicate the intake of these total nutrient compounds that the bird needs in order to be healthy and to give us good breeding results.

The ways we can effectively duplicate the natural habitat nutrients are to give good and complete grain mixes and various oils, and to use chemical, vitamin or nutrient additives, fruits, fruit supplements, soft food mixes and sprouted seeds. It would seem that, at least from my experience, the primary supplemental items that the birds want are apple and soft food mixes, including sprouted seeds and greens. They seem to have a time clock in their heads. If you don’t give them the mix they are used to on a given day and/or you don’t give it to them at approximately the time of day they are used to getting it, they certainly are vociferous in letting you know about it.

One of the primary supplements given near and during breeding season is fresh corn on the cob. The birds seem to enjoy it, it is nutritious, is easy for the hens to make crop milk, and thus takes quite a strain off of the hens in feeding the young.

Another type of feeding that we have on occasion employed or will encounter is Gevage, hand feeding, tube feeding or whatever you may call it. This method of feeding may be done by choice or by necessity. There are on occasion hens that will refuse to feed their young or may feed them to a certain point and then discontinue feeding. Rather than take a chance that the young will perish in the nest with a bird of this type, we naturally will pull them and hand feed. We can further increase the productivity of these birds by permitting the to lay and hatch their eggs and then pull the young and hand feed them. We generally find that these hens will go back to the nest, though we only permit them to have one further clutch as we do not feel that over-breeding can be of any advantage to the bird.

We know that we cannot duplicate the exact foods that the parent birds would ordinarily feed their young. However, we do attempt to come as close as possible by making up mixes of grain and cereals with supplemental additives, such as vitamins and electrolytes. We cook these mixes and thus give it some semblance of the pre-digested quality that the parent will have when feeding the baby. The formula that we use is a fairly standard one, made up basically of safflower or soy meals, baby foods of high protein level, oatmeal, cream of rice, wheat heart cereal, masa harina (which is a corn meal) with additives such as baby vegetables (predominantly Spinach) for the greens, tri-calcium phosphate to help the growth of the babies’ bone structure and vitamins. Through experience we have found that this particular formula is successful in feeding baby birds literally from day of hatch to the day they are weaned. We have used it successfully on every Psittacine bird we have in our collection with the exception of the pure fruit and nectar eaters, such as lories.

Another key area of consideration in breeding of psittacine birds is housing. These do not have to be aviaries of any particular type, size or method of construction. We experienced successful results in breeding many birds, including some of the larger parakeets, parrots and macaws, in relatively small breeding cages. I am quite convinced, however, that nutrition is the primary factor in bringing any bird

into good condition and ultimately breeding that bird. However, when we do breed birds in small cages, we give those birds rest periods in larger flight areas so that they can maintain their good body condition through exercise and normal flight. Some of the largest birds, such as macaws and cockatoos, have been successfully raised in amazingly small cages by many aviculturists. Macaws have been known to regularly breed in areas as small as two and one-half feet by three feet. The breeding urges are rather strong in all these birds, and unusually confining conditions do not necessarily preclude successful breeding. One aspect we have found that will at times preclude breeding, particularly in cockatoos, is where a bird has been hand fed and played with and made inordinately tame. We do find many of these birds will have absolutely nothing to do with other birds, and we have found no practical way to get them to breed. When we hand feed babies we want to use as breeding stock, we handle them only for the short period of time necessary to get the food into them and then ignore them, leaving them (hopefully with other babies) in the nest or box in which they are held during feeding. We find that this gives these birds a far greater opportunity to realize they are birds.

An important feature in any aviary or cage is the perches. Both the diameter and the stability or steadiness of the perch are of paramount importance. Naturally, if the perch moves, proper mating frequently cannot take place. Further, the comfort of the birds and their ability to hang on to the perch, particularly the hens, is of great importance. The perch should be of such size that when the bird has its feet clamped on it, the claws are approximately two-thirds of the way around the total circumference of the perch. One of the best types of perches is the natural tree limb. If possible, stay away from the dowel perch, which is often found in commercial cages and, I am afraid, all too frequently used by breeders. The dowel is much too round, smooth, small and, in general, unnatural for the bird. To find a dowel of adequate size in a hard wood that these birds will not eat up in a matter of minutes is very difficult. Realizing that all psittacines are chewing animals (as they need this chewing to keep their beaks in good condition), we certainly should give them something to chew on, hopefully something other than their perches. Assuming that we use perch woods hard enough that the birds cannot destroy, such as dry eucalyptus, then provide some other wood, such as 2 x 4’s in small pieces, hung in various places around the aviary so that the birds will have something not only to keep them out of mischief but to give them the constant ability to satisfy their instinctive chewing action.

Nest boxes also play an important part in the breeding of psittacine birds. There is no such thing as an absolute rule in the use of a nest box of given size or type for any particular bird. We have seen a pair of birds use a box 9 x 9″ x 6′ deep for their first clutch and for the second clutch in the same aviary in the same year use a box that is 10 x 12″ x 12″ deep. However, we do attempt to standardize on a particular box for a given type of bird. We have found in approximately eighty percent of our breeding stock that the same size and type of box will be readily accepted year in and year out by the same species of birds. When we find birds that do not seem to be well settled or properly acclimated to a particular box, these birds will be given a choice of up to four different types, sizes and shapes of nest boxes. We will let them pick the box they want, and if they seem totally content in that box, we will pull the others. On occasion we will leave the other boxes simply because the birds do not seem to be fully content, and we generally find that in subsequent clutches they will take a different box. As a general rule, we attempt to make our nest boxes approximately one to one and a half inches greater in width and front-to-back dimensions than the bird is long from head to tail. The only exceptions to this are those rather long tailed parakeets, such as the Ring Neck, Princess, Plum Head, etc. In those cases we found that a box approximately one to two inches longer than the body of the birds seemed not only to be satisfactory but generally preferred by the birds. In the case of large macaws, we use steel drums or garbage cans, generally the thirty-two gallon size, and the various cockatoos will use anything from twenty gallon steel drums up to fifty-five gallon steel drums.

Of paramount importance in any nest box is a ladder which the bird must have. Otherwise, many hens will enter the nest box by jumping on the eggs, subsequently breaking them and/or killing young that may be in the box. These ladders should be made of wire and preferably spaced away from the inside frontal facing of the box at least one quarter to one half inch so the birds cannot get their feet or nails hung up.

The hole for entry is generally ten to fifteen percent wider than the dimension of the body of the particular bird to use that nest. There are certain exceptions to this, such as in the case of the Blue Bonnets where the hens are obviously much smaller than the males. In those cases, we make a box where the body of the bird will easily fit through the hole and then cover it entirely with a piece of bark, drilling only a small eighth inch or quarter inch hole. The hen will pick out her own entry to the exact size of her own body, thus precluding entry to the male. He, for some reason, will not attempt to get into the box. However, should you leave a normal nest entry hole, you will ordinarily find that the male will go into the box, disturbing the hen and frequently killing the young.

Other types of next boxes used for various birds can be found of a specialty type. A good example would be Hooded Parakeets where we found successful breeding in two types of conditions. One is a bale of moss where the birds will burrow into the moss, generally from the top, going almost straight down and then off at an angle. We also make nest boxes for these birds in an “L” shape where they can enter the box, go down and off to the side. We found with these particular birds (evidently being termite hill or ant hill nesters in their native habitat) that they have a peculiar habit of circling the box. As a result, we staple bark on the outside of the box, not for a natural appearance but merely to give them a footing. We find that the hens prefer to be able to run entirely around their boxes.

I realize that many people refer to any box of more than two feet in depth as being a grandfather type. However, the majority of our nest boxes range in depth from sixteen to thirty inches. What we consider the grandfather boxes are nests that are from five to seven feet in depth. Particular birds that seem to enjoy boxes of that type are the King Parakeets, Crimson Parakeets, Barraband Parakeets and the Princess of Wales. (The last bird mentioned is one I have seen very successfully breeding in boxes that are normally construed as cockatiel boxes.) As a generality, it is wise to make the size of the nest box smaller, rather than larger, than your estimation of what truly is necessary.

We had some problems several years ago in attempting to breed Plum Head and Blossom Head Parakeets. We found them constantly chewing through the bottom of the box and invariably cowering in one corner. Though we occasionally had clutches of eggs, we never really, in our estimation, successfully bred these birds. I finally designed a box that has the shape of an “A” frame where the top of the box is very narrow, only about three and a half inches to four inches in width. The bottom of the box is also quite narrow, approximately eight inches in width and nine inches front to back. These boxes are thirty-six inches deep. Since that time we found the Plum Heads are very successful breeders, all of them preferring this type of box.

The nesting material which we place in the boxes is fairly standardized for most birds. We happen to be fortunate in that we have a ready supply available to us of coarse shavings (not sawdust, which we know can be dangerous as babies can ingest the material and die from it). We mix the coarse shavings approximately fifty percent with rotted wood and bark which has been sterilized by placing it in an oven at approximately 200 degrees F for a period of forty-five minutes to one hour. Once this mix is made, we will add one-third sterilized potting soil, which can be obtained from any nursery, and two-thirds of the previous mix. We then make this quite wet, squeeze all the excess moisture out and load the boxes from one and one-half to three inches deep with the mixture. We use a sterilized potting soil for two reasons, (1) because it retains a substantial amount of moisture as it has various types of moss and humus in it and (2) it is sterilized and thus precludes the possibility of unwanted and dangerous bacteria which other types of dirt frequently harbor.

If we are fortunate enough to select our breeding stock and pick only the best breeders, we certainly have precluded many of our problems. However, the majority of us are not so fortunate. As a result, we occasionally find hens that are going to jump into their nest boxes, destroying the eggs and/or the young, regardless of the ladders we may install. I have seen specialty boxes constructed to curtail this wherein a long tunnel was built extending from the hole of the box. The bird lands at the entrance of the tunnel and has to walk at a rather slow pace to get through the narrow tunnel until she gets to the nest box itself. We have not had much success in using this. However, I have seen other aviculturists use it quite success- fully. We use another method to curtail the problem. We create a cross of perch material that will be installed inside the nest box, going from front to back and side to side, generally installed approximately two to three inches above the height of a sitting hen in the bottom of the box. We discovered that the hens will jump onto these cross perches and then rather gingerly step down to the bottom of the box. This has eliminated a lot of problems of that type.
Behavioral patterns are of great import in the successful breeding of psittacines. However, few of us really have the opportunity to observe our birds to learn all their habits. The peculiarities of each bird can be as varied as the differences found in human beings.

We find in certain birds, particularly the rosellas, that the hens seem to come into breeding condition slightly after the males. Thus, the males are quite active in attempting to breed and force the hens toward the nest boxes, driving them quite hard on occasion. Death sometimes results. To preclude this possibility, we catch the males and cut every other primary feather back to the point of the secondaries on one wing. They can still fly, yet it slows them down enough so that the hens can stay ahead of the males. On one occasion approximately three years ago a pair of Golden Mantels that had bred successfully for approximately five years went into the normal breeding season. Since I knew the bird well, I did not observe them too closely. What I did not realize was that the hen evidently came into breeding condition earlier than the male. The female, acting as the aggressor, debeaked the male, and we lost him. If I had not seen it, I would not have believed it. This particular case illustrates the fact that we can never know our birds too well. Constant observation is always necessary for successful breeding. It is very important to get to know on a personally familiar basis every bird in our breeding collection. It is not realistic to expect all individuals of a species to follow the exact same pattern. We must expect variances from bird to bird.

The proximity of birds to each other is also important. We attempt to keep pairs of the same species separated by at least three or four flights. All flights should be double-wired because birds are likely to fight with adjacent neighbors when they are in breeding condition. There are certain exceptions, however. Pileated Parakeets, for example, have far greater breeding success when we keep them in every other flight. It seems when one male comes into breeding condition, each male down the line follows his lead. When widely separated, breeding is spotty and scattered. We have also found greater success in keeping the Princess of Wales Parakeets close enough to hear each other. The call of one bird tends to bring the others of the same species into breeding condition. On the other hand, we never put two varieties of rosellas next door to each other.

There are a few psittacines that can be successfully bred in colonies rather than in individual pairs. The love birds fall into this category with the exception of the more difficult species such as Abyssinians and Madagascars. Budgerigars certainly will colony breed, and this is often done when breeding for commercial quantity rather than for quality. We find, however, only one Australian parakeet, the Crimson Wing, is distinctly more likely to breed successfully in colonies than as single pairs. All cockatoos will also colony breed quite successfully. The key point with the cockatoos is to be certain that the nest boxes are widely separated, preferably at either end of the flight. As long as the birds can stay out of the way of each other, they seem to do better when they have company in their flight rather than one pair per flight. With cockatoos the size of the flight cage is not really of great importance.
I am firmly convinced that virtually any psittacine can be cage bred, including birds as large as cockatoos and macaws. All of our flights are the same size. They are four feet wide, fifteen feet long and eight feet high at peak of the roof. We have a drop front on these flights, thus making them draft free. Roosting perches must be in an area that is draft free. The wire areas of our flights are eight feet long. They are approximately eighty inches in height, extending out from front. The rear of the flight is approximately seven feet long and entirely made of plywood. In certain of our flights, such as those for cockatoos and macaws, we wire the entire interior of the plywood area so they cannot chew their way out.

Temperature in the aviary (and particularly the chill factor in cold weather) will be warner in the roosting area. We have not actually taken the temperature of the birds. However, it is known that many animals tend to lower their body temperature when they sleep. We believe this may well be the case with the birds. Since birds have normally high metabolism rate and body temperatures, we believe that it is important that they be given as much shelter as possible when roosting. One of the greatest detriments to our Dirds is drafts. A draft does not necessarily mean cold air. Moving air constitutes a draft for a bird. Many times we have seen pet birds kept indoors at absolute constant temperatures. Then the slightest draft, such as a forced air furnace going on and creating moving air, seems to lead to upper respiratory infections or so-called “colds”. Cold air tends to go in straight lines, not around corners, and if an aviary is properly constructed, the birds will be able to remain totally draft free.

The directions the aviaries face are not nearly as important as has been reported on occasion in the past. The primary or preferred direction in the northern hemisphere has been stated as south, the second most preferred direction is east, etc. Our aviaries face virtually every direction, and we found great success in breeding our birds in any one of these aviaries. Should we find a pair of birds that does not breed in a particular aviary, we may move them to another cage, perhaps to an aviary immediately adjacent to the one they were in or perhaps at the opposite end of the breeding pens.

In-breeding any birds can be disastrous. This should never be done because it can ruin the strain, strength and, ultimately, the overall health of the birds. We know that when birds (particularly budgerigars, are bred for show purposes, in-breeding and line-breeding are used. Line-breeding is slightly varied from in-breeding in that there is (to a degree) a certain amount of out-crossing of stock. However, my personal belief is that line-breeding is just a sophisticated way of saying in-breeding. The attempt to get the best show animal and the best pedigree line frequently seems to cause some of the weakest animals and those must susceptible to sickness and disease. We know that certain faults can be eliminated in birds through selective theoretic line-breeding. However, what constitutes a fault to one person may not necessarily be a fault to another. The so-called show standards which have been set up for birds seem to vary from show to show and from judge to judge except for budgerigars.

If this happens to be your forte, do what you see fit in the breeding of your birds, but remember first, foremost and always, that the strength of the bird is far more important to continual successful breeding than the appearance of its shape or color.

In summary, we cannot say that all birds are bred in captivity the same way. However, in general, we can say that the majority of psittacines are effectively bred the same way with few exceptions. First, nutrition is of greatest importance. Second, proper surrounding or housing is essential, whether it be cage or aviary. Third, adequate and properly designed nest boxes, as to size, overall dimension, depth and nesting materials, is important. Virtually any psittacine will breed within a reasonable period of time upon proper acclimation to its surroundings, its conditions and the aforementioned necessities. There are no mysteries in the breeding of these birds, only the application of good common sense with judicious observation of your birds and a few simple rules properly applied.

BEHAVIOR GENETICS STUDIES WITH BIRDS

By Luis F. Baptista

IFCB 1978

INTRODUCTION
Each animal possesses a genome which endows it with dispositions to respond in a certain range of ways, to a certain range of appropriate stimuli, and thus feeds itself, finds shelter, acquires a mate and contributes to the gene pool (Manning 1972). Gene mutations are continually emerging. As environment changes, genomes may also have to change to adapt to these changes. Thus, individuals possessing dispositions to perform new behavior patterns that enable them to cope with environmental changes will be selected for and leave progeny. The heritability of behavioral traits has been a long debated issue often referred to as the “Nature versus Nurture” controversy. Ethologists have long been aware of the heritability of behavior patterns (nature), whereas many psychologists, preoccupied with learning phenomena (nurture) were slow to be convinced.

That dispositions to perform specific behavior patterns may actually be inherited and thus be subjected to the forces of natural selection, comes from studies of interspecific hybrids, artificial selection of extreme behavior traits, and intraspecific hybrids between selected strains of animals. Data from studies on invertebrates are plentiful (Gould 1974). Data on birds are relatively few. This paper reviews some of the changes brought about during the process of domestication of birds and summarizes studies on behavior of natural and artificial interspecific avian hybrids. These include my own unpublished studies on hybrid Estrildid Finches study of domesticated species can be very instructive as to how natural selection operates on behavior, especially if the ancestral forms are still available for comparison. Both natural and artificial selection (e.g. domestication) operate on mutations resulting in reduction of variation (Frank 1974). The process of speciation requires isolation of gene pools. Domestication is the result of artificial isolation of segments of a gene pool from the wild population (Goodwin 1965, Nicolai 1976).

EFFECTS OF DOMESTICATION

Tameness

Canaries
Birds that lend themselves to domestication must be easily maintained and bred in captivity. Indeed, Goodwin (1965) points out that most domesticated forms are granivorous species (Galliformes, Columbiformes, Psittaciformes, Fringillids and Estrildids), the groups that are easiest to maintain in captivity.

Man must also have selected for individuals that would breed readily in captivity. Throughout the world today, societies exist for the propagation and perpetuation of the many domesticated breeds of canaries (Serinus Canaria). These birds are docile and are often bred in small cages. Nicolas (1960) compared the behavior of domesticated and wild caught canaries. He found that wild birds remained skittish for a long time while held captive and seldom came into breeding condition. Clearly, man must have selected good breeders as ancestors of our domesticated forms. That tameness in domesticated canaries is heritable was demonstrated by Hinde (1956). He presented canaries, Goldfinches (Carduelis carduelis) and hybrid Goldfinch x canaries with stuffed owls. Goldfinches mobbed the owls, but many canaries showed only a passing interest in the models. The latter was interpreted as a by-product of artificial selection for tameness by man. Hybrid Goldfinch x canaries also mobbed the models but less vigorously than pure Goldfinches.

Turkeys
Loepold (1944) studied the heritability of tameness in turkeys (Meleagris gallopavo). Wild turkeys are difficult to keep and breed it captivity, frequently fighting so that game farms usually cross domesticated birds with wild ones and release the hybrids for the hunter’s gun. Leopold noted a lack of alertness and wariness in wild hybrids. Wild birds had an uncanny ability to see the observer before he spotted the birds and would immediately run for cover. hybrids could be approached more closely. If the observer withdraw a few hundred yards at the first sign of alarm from the hybrids, the latter would resume normal activity and permit further observation.

Ducks
Another measure of tameness is the speed at which an animal habituates to a novel stimulus. Desforges and Wood-Gush (1975a) presented wild mallards (Anas platyrhynchos) and domestic Aylesbury Ducks with a model of a stuffed brown Leghorn Hen. The domestic breed habituated faster to the presence of the model. They then replaced their food pellets with colored pellets. Although the latency to respond increased in the Aylesbury, they eventually consumed the pellets. The mallards never ate the colored pellets.

Recognition of Young

Chickens
Heinroth (in Lorenz 1971) noted that females of Burmese Jungle Fowl (Gallus gallus) will only lead chicks with markings on the head typical of the wild form. Chicks of domesticated breeds lacking head markings will be killed. This recognition of species’ specific characteristics in chicks is still found in some domestic breeds (e.g. some gamecock breeds, Phoenix chickens). However, in many domesticated breeds man has apparently selected against the ability to recognize species’ specific characteristics so that even ducklings or goslings may be brooded and led. Indeed, in some breeds (Orpington, Plymouth Rock) even the response to the acoustic character of the releasing mechanism has been selected out so that hens will even try to care for young mammals (Lorenz 1971).

Estrildid Finches
Nestlings of Estrildid Finches have species’ specific colors and markings on their palates and beg with species’ specific head-twisting movements and vocalizations. That these palate markings serve as species’ specific releasers was tested quantitatively by Immelmann, et al (1977). The authors placed nestlings of domesticated strains of Zebra Finches (Poephila guttata) which lacked palate markings with “wild-colored” Zebra Finch nestlings which had five black spots on their palates. Wild-colored nestlings had significantly higher survival rates, had priority to the first feedings of the day and received more food during the later phases of the nestling stage so that they grew at a more rapid rate. Nicolai (1964) performed cross-fostering experiments with various Estrildid species. He noted that these finches would not feed newly-hatched chicks with mouth markings differing from those of their young. If mouth markings of fostered heterospecific nestlings were similar to those of their own, the strangers would be fed for a while and then deserted.

As in some domesticated breeds of chickens, man has bred out the selectivity of response to the species’ specific releasers (mouth markings) of their young in the domesticated Bengalese Finch (Lonchura striata). Guttinger and Nicolai (1973) report that Bengalese Finches in their experiments raised 122 individuals belonging to 15 species in 7 genera (Lagonosticta, Uraeginthus, Ptyilia, Spermestes, Lonchura, Heteromunia, Erythrura) with diverse palate markings. In other cross-fostering experiments, Bengalese have been used to raise the genera Poephila, Euodice, Chloebia and Odontospiza (Immelmann 1969, Baptista 1973a, 1973b). Indeed, Gouldian Finches (Chloebia gouldiae) are frequently raised by Bengalese Finches in Japan and exported to Europe (Nicolai 1967). The list of genera raised by Bengalese is probably incomplete.

Social Behavior

Turkeys
Leopold (1944) found differences in the social behavior of wild turkeys and free-ranging wild x domestic turkey hybrids. Wild turkeys tended to gather in small flocks ranging from 2 to 12 animals (X 2.5 to 6.3 between 1940 and 1943). Even at low densities, hybrid turkeys gathered in large flocks ranging from 7 to 33 birds (X 14.8
to 19.2 between 1940 and 1943). There is a partial segregation of the sexes in wild turkey flocks, i.e. adult males tended to winter separately from adult females and young of the year. This segregation was not found in the hybrid strains.

Pigeons
Edrich and Keeton (1977) performed homing experiments with feral and domestic “homer” pigeons (Columba Livia). Ferals had on the average longer homing times and lower return rates than “homers”. This was partly due to weaker homing drives in the feral pigeons but partly due to a selected change in the social behavior of homers. Feral pigeons tend to be easily “distracted” by other pigeons, tending to join other pigeons flocking in the air or on the ground. Man has apparently selected out this tendency to flock in the homer.

Cockerels
A number of game-cock breeds have been selected for aggression. Fennel (1945) compared the behavior of some game-cock breeds and more docile domestic chickens. He found that game-cocks were able to tolerate more punishment, were shiftier, faster and not as clumsy as “domestic” cocks. He also noted that different breeds utilized different methods of fighting. “Kentucky Dominiques” attack from above by flying at the opponent. “Allen Roundheads” move to within striking distance, then strike forward or toward the enemy on either side.
Cockerels have also been selected for high and low sex drive through three generations (Wood-Gush 1960). Siegel (1965) performed bidirectional selection experiments for completed matings over six generations.

Japanese Quail
Kovach (1974) has reviewed the literature on genetic factors in sexual behavior in this species. Strain differences have been found in the sexual receptivity of females. Another investigator successfully undertook bidirectional selection for high and low mating activities in male quail in one generation. Strain differences were also found in mating frequencies.

Ducks
Desforges and Wood-Gush (1975b) reported selection against aggression in domestic Aylesbury Ducks. Individual distance was smaller for Aylesbury than for mallards during feeding and resting. For example, mallard drakes maintained an individual distance of
45.7 cm from another drake during feeding, whereas the domestic drakes maintained distances of only 30 cm.

Vocalizations

Domestic Chickens
Kruijt (1964) studied the ontogeny of behavior in Burmese Jungle Fowl (Gallus gallus), ancestor of our domestic breeds of chickens. When distressed (hungry, cold or isolated), chicks utter a loud shrill call at the rate of 2 to 3 per second. Kruijt’s spectrograms showed that shrill calls of domestic chicks differed in morphology from those of their wild ancestor.

Konishi (1963) showed that crowing in domestic cocks is genetically determined, developing normally even in birds deafened at the time of hatching. Selection for differences in crow duration in two different breeds was demonstrated by Siegel et al (1964) who studied Athens-Canadian Random breds and White Rock roosters. Crows of White Rocks were significantly longer. They suggested polygenic inheritance in crow duration as evidenced by the fact that variation between individuals was greater than within individuals

Crow duration in the above breeds averaged 1.58 to 2.43 seconds. Masui (in Siegel et al 1964) reported that the Totenko breed of ornamental chickens had crows lasting 15 to 20 seconds, again a result of artificial selection.

Goodwin (1965) noted that males of most of the largest breeds of domestic chickens have crows that are more long-drawn with the last syllable lengthened out, whereas bantams have shorter crows more similar to the ancestral Gallus gallus.

Domestic Ducks
When performing the sexual display known as the grunt-whistle (see Figure 2), mallards utter a clear flute-like whistle. Desforges and Wood-Gush (1976) found that in one of the domestic breeds (the Aylesbury) the whistle has been reduced to a low grunting sound.

Domestic Pigeons
In his treatise on domestic animals, Darwin (1875) pointed out that man has selected for distinct qualities of voice in at least two breeds of pigeons. Levi (1965) describes 13 “voice” breeds known collectively as “Trumpeters). A second voice breed noted by Darwin is known as the “Laugher.”

behaviorgenetics_pict222

Figure 1. Principle components of distress calls of pheasant, chicken, small hybrid (H1) and large hybrid (H2). After McGrath et al. 1972.
behaviorgenetics_pict223
Figure 2.
First row, left to right:
1. Mallard, grunt-whistle
2. Mallard, headup-tailup
3. Mallard, downup
4. Mallard, nod-swim

Second row, left to right:
1. Pintail, grunt-whistle
2. Pintail, headup-tailup
3. Pintail, burp

Third row
Mallard, bridling.

After Lorenz, 1971.

Levi (1965) describes the courtship song of the Thailand Laugher as a “who-a, who-a” followed by 8 or 10 wock-wock, wockwock. The male English Trumpeter begins its courtship song with several coo-coo-roo-coo’s similar to the ancestral Rock Dove, but this is followed by a long series of who-oo-oo-oo-oo-oo, who-oo-oo-oo-oo-oo, etc. (Baptista and Abs 1981). The peculiar sounds used by other breeds are also used in the nest showing ceremony. In this display the male sits in the nest cup, bows rhythmically, flicks the wings up and down rapidly and vocalizes. Another breed, the Altenburg Trumpeter, has a much more rapidly uttered series of vocalizations following the courtship coo. The Altenburg’s trumpeting is also more uniform in rhythm than in the English breed (Baptista and Abs 1981).

Domestic Quail
The Japanese Quail (Coturnix coturnix japonica) was domesticated as a “good song bird” during the Muromachi era, some 600 years ago (Yamashina 1961). In Central Europe, Coturnix Quail were independently selected for qualities of voice. All breeds of Japanese singing quail were lost during World War II, and the European strains were lost because of loss of popularity of the hobby (Howes 1964).

Goodwin (1965) noted that wild Japanese Quail give a call (crow) which sounds like “Quah–kah”. Domesticated birds give a call which sounds like “Quah-grrr”. This was selected for because it suggested to the Japanese the sound of “distant thunder”. During the Tokugawa Period, they were kept in elaborate cages and judged in competitions on quality of song. Moreau and Wayre (1968) compared sound spectrograms of wild Japanese Quail with extant domestic stock. Although calls of wild quail are similar to those of the domestic birds, the latter are “louder” and harsher.

Greater and Lesser Prairie Chickens
The vocalization associated with the “booming” display in a hybrid consisted of six syllables in contrast to the three syllables in both parental species. Duration of the directed display was 1.96 seconds in the greater, 0.60 seconds in the lesser, versus 0.97 seconds in the hybrid (Crawford 1978).

Canary
Along with differences of appearance in the various domesticated canary breeds, man has selected for different qualities of voice/ Marley (1959) studied spectrograms of Roller Canary songs and compared them with songs of their wild ancestors. Roller songs are softer, lower pitched, consist of long repetitive trills with shorter intervals between notes and are simpler in structure than songs of wild canaries. Poulson (1959) also noted that Rollers sing with their bills virtually closed, whereas “common” canaries (= choppers?) sing with their bills open. Common canaries may imitate Roller songs but sing them with bill open. He found also that the soft and low pitched quality of the songs developed in Rollers raised in isolation. Syllable diversity, however, is learned (Poulson 1959, Marler and Waser 1977, Waser and Marley 1977).
Poulson (1959) also compared two social calls of Roller and “common” canaries. Both contact and alarm calls of the Roller Canary are also lower pitched than those of common canaries and exhibited differences in patterns of pitch modulation. Recent studies by Murdinger (1970) indicate that at least some social calls in Carduelid Finches are learned. Thus the relative contributions of nature versus nurture in the development of canary calls remains to be ascertained.

Courtship

Pigeons
As a result of artificial selection by man, various breeds of domestic pigeons exhibit breed specific behavior patterns. The latter appear to be derived from various components of courtship and other social displays of Columba livia.

Pouting
Columba livia inflates its neck with air (pouts) during courtship and during the nest-calling display. Pouting may continue for a while after courtship (Whitman 1919). In the 40 or so extant breeds of pouter pigeons, pouting is exaggerated and occurs at the slightest disturbance, e.g. during aggressive displays against conspecifics or a human intruder. Pouting is, therefore, the hypertrophy of a universal instinct in pigeons to inflate the neck (Whitman 1919, Nicolai 1976).

Wing Lifting
The aerial display of C. livia is territorial and sexual in motivation. In this display the pigeon claps its wings loudly several times, then glides with its wings held slightly above the horizontal, finally gliding with its wings held out horizontally. The Swing Pouter holds it wings almost vertically above its back during the gliding phase so that it quickly loses altitude and must flap its wings to return to a greater height.

Wing Clapping
Two breeds, the Swing Pouter and Rhine Ringbeater, perform exaggerated loud clapping phases during their display flights. This leads to premature fraying of the primaries early in the breeding season. The clapping phase of the flight display is lengthened in the Swing Pouter. Nicolai (1976) once counted as many as 30 claps during the display flight of this breed, whereas it is usually 4 to 6 claps in the ancestral C. Livia. Nicolai crossed these two loud clapping breeds with some of the normally (softer) clapping breeds, and found that loudness was intermediate in the Fl generation.

Rolling
Levi (1965) has cataloged some 22 breeds of roller and tumbler pigeons. These may be further segregated into flying rollers, parlor rollers, flying tumblers and parlor tumblers. Tumblers differ from rollers in that whereas tumblers perform only one or two backward somersaults per series, rollers may perform many, often falling 50 or more feet in the course of rolling. Young individuals of both types fly normally for the first weeks or months but eventually all perform intermittent somersaults. The parlor types eventually lose their ability to fly altogether, rolling or tumbling on the ground.

Nicolai (1976) traced rolling behavior in Birmingham Rollers to the glide phase of the display flight, noting that loss of altitude is a prelude to rolling in young birds. However, Entrikin and Erway (1972) and Kerry Muller (personal communication) report that any attempts to fly initiates rolling or tumbling in the air or on the ground.

Nicolai (1976) crossed rollers with non-rolling breeds and noted that Fl hybrids could perform only incomplete somersaults. A more detailed study of the inheritance of rolling behavior in pigeons was performed by Entrikin and Erway (1972). Crosses between flying rollers and normal breeds produced 38 non-performers and one bird that somersaulted only once or twice. F2’s were split 50-50 into rolling and non-rolling birds. F1’s back-crossed to flying rollers produced about equal numbers of performers and non-performers. Flying rollers crossed with parlor rollers produced about equal numbers of parlor and flying rollers. The authors suggest that rolling behavior is controlled by a single autosomal mutation. However, the penetrance and expressivity of the gene appears to De modified by other genetic factors.

Ringbeating
During its ground courtship bow-cooing display, a domestic pigeon may sometimes jump into the air, clap its wings loudly several times, then drop just behind or beside the female and recommence his bow-coo display. In the extinct English breed called the Finnikin (Goodwin 1965) and the extant German Ringbeater (Nicolai 1976), the clap display is elaborated so that the courting male will fly one to four times in a tight circle over the female with a constant loud clapping of the wings. Nicolai points out that even the female Ringbeater claps louder than other breeds, although not as loudly as the male.

Nicolai isolated an unpaired female Ringbeater for some time, and then placed her in the middle compartment (1.5 x 2.5 x 2.3 m) of a three-compartment enclosure. In one of the other compartments he placed a male Ringbeater and in the other he placed a male of another breed. She was visually isolated from both males by partitions of cloth but could hear both displaying. She was attracted to the clapping display of the male Ringbeater and ignored the other male. The author suggested that a parallel change in the signal receiver also occurred which was genetically fixed with artificial selection.

Ducks
Desforges and Wood-Gush (1976) compared the courtship displays of domesticated Aylesbury Ducks and the wild ancestors, the mallard. As mentioned earlier, the acoustical component in the grunt-whistle display of the Aylesbury has been reduced to a grunt. Following the head-up-tail-up display, mallards turn the back of the head to the female. The last display is absent in the Aylesbury. Moreover, the preliminary shaking preceding the head-up-tail-up display of the mallard was sometimes absent in the Aylesbury.

The down-up display was performed at lower intensity by the Aylesbury. The nod-swim display was less often seen in the female Aylesbury and reduced to an intention movement.

Pair-formation displays were always directed at only one female in the mallard. Aylesburies would direct these same displays (e.g. precopulatory pumping, preen-behind-wing) at several females. Mallards were thus thought to form pairs in captivity, whereas Aylesburies did not.

Behavior of Hybrids

Maintenance Activities

Diving versus Dabbling in Ducks
The different tribes of ducks tend to forage in different ways. Mergansers (Mergini) dive beneath the water’s surface to pursue their prey. Shelducks (Tadorna tadorna) belong to the surface duck tribe (Anatini) which typically feed by (1) “dabbling” on the water’s surface to pick up vegetation and small animals in their bills or (2) “tipping”, wherein half the body is submerged under water, and the hindquarters remain on the surface. Lind and Poulson (1963) studied the feeding behavior of a Goosander (Mergus merganser) x Shelduck hybrid. The latter never dived but dabbled and tipped like its Shelduck parent. However, whereas Shelducks tip with their body 90° with the horizontal, the hybrid tipped at 45°. The authors attributed tipping angle to the body shape of the hybrid.

Scratching versus Digging
Typical of many galliformes, Lady Amherst Pheasants (Chrysolophus amherstiae) feed by scratching the ground with alternate backward movements of the feet. Impeyan Pheasants (Lophophorus impeyanus) dig the soil with the bill when feeding. A hybrid Impeyan x Lady Amherst Pheasant fed by digging with its bill like its male parent (Huxley 1941).

Foraging for Worms
Whitman (1919) observed that whereas many species of pigeons and doves feed occasionally on earthworms, Passenger Pigeons (Ectopistes mi_gratorius) seemed to relish these food items more than did other species. A Passenger Pigeon hatched under Ring Doves (Streptopelia risoria), never having seen an earthworm before, immediately approached the worms when presented with them, handled and ate them. Whitman went on to present one naive Passenger Pigeon x Ring Dove hybrid and two “pure” Ring Doves with some earthworms and soil. Whereas the Ring Doves ignored the earthworms, the hybrid immediately approached the worms and quickly acquired the skill of extracting worms from their burrows and eating them. Whitman suggested that the recognition of earthworms as preferred food was inherited from the Passenger Pigeon parent.

Holding Items with the Foot
A number of avian taxa have independently derived the ability to manipulate food items or nest material with the feet (Clark 1973). Even within the same family or subfamily, species differ in the frequency with which this behavior is exhibited. The European Goldfinch frequently uses its foot, the canary sometimes and the Greenfinch (Carduelis chloris) rarely. Hinde (1956) noted that several Goldfinch x Greenfinch and Goldfinch x canary hybrids often used their feet, whereas Greenfinch x canary hybrids seldom did. Hybrids inherited their holding ability from their Goldfinch parents.

Among Estrildid Finches, Zebra Finches never use their feet to hold items on a perch, whereas African Silverbills (Euodice cantans) regularly do so. Four Silverbill x Zebra Finch hybrids regularly manipulated seeding heads of grass with their feet, indicating inheritance of a dominant trait (Baptista personal observation).

Biting Versus Pulling of Grass
Poulson (1950) observed that Greylag Geese (Anser anser) are equipped with “tooth-like knots” along the sides of their mandibles enabling them to “bite” grass while grazing. Ducks are not so equipped and thus “pull” grass while foraging thereon. A hybrid domestic duck x Greylag Goose bit grass like the goose parent. However, not being equipped with the bill “teeth” of the goose, the hybrid bit grass with the tip of the bill which was equipped with a large nail.

Drinking Behavior
Most finches in the family Estrildidae drink in the manner of chickens, i.e., water is scooped into the bill, and the head is subsequently raised so that the bill is pointing skyward and the water runs down the throat through gravity. Some Australian Finches in the genus Poephila, independently of the doves and pigeons (Columbiformes), drink by dipping the bill in the water and sucking without raising the head. Immelmann (1976 and personal communication) studied drinking behavior in hybrids between Mannikins (Lonchurae) which scoop, and Poephila species which suck. Hybrids sucked in pigeon fashion, but bills were not dipped as deeply into the water as in Poephila species.

Vocalizations

Chickens x Pheasants
McGrath et al (1972) artifically inseminated female Ring-neck Pheasants (Phasianus colchicus) with semen from White-Leghorn Chickens. One year old hybrids were of two size classes: males weighted 1,775 gm and females weighed 875 gm. Adults of both parental types and their hybrids were hand-held with the wings and their distress calls recorded.
Duration and pitch of the principal component (= darkest and most distinct harmonic) of these calls were measured (Figure 1). Duration of hybrid calls was similar to that of the pheasant. Pitch of the hybrids was intermediate between the parental forms. However, the smaller hybrids had higher pitched calls than the larger hybrids. There was little change in frequency in the chicken calls, whereas there was an initial rise and then a fall in pitch in the pheasant calls and those of the hybrids. Harmonic structure of some of the hybrids of both size classes resembled either the pheasant or the chicken.

It was suggested that the inheritance of duration was controlled by one or few genes, whereas principal component (pitch) was polygenic.

Palearctic Quail
Moreau and Wayre (1968) studied the “crows” of European and Japanese “races” (species?) of Coturnix coturnix. The call of the European bird was clear, brief and consisted of sharp notes, whereas Japanese Quail calls were harsh and blurred on the spectrograms and were less far-carrying. Moreover, the rhythm of the European’s call was “dactylic”.
Calls of European x Japanese hybrids approximated the European’s in rhythm but showed a coarsening in the second and third notes that characterized Japanese calls. Calls of Japanese x European hybrids showed nothing approaching the European rhythm and only a limited coarseness. Although more or less intermediate, hybrid calls tended to be more similar to the male parent.

Greater x Lesser Prairie Chickens
Courtship displays of Prairie Chickens (Tympanuchus spp.) consist of drooping of wings, tail-fanning, erection of pinnae, inflation of vocal sacs and the “booming” vocalizations (Crawford 1978). The Greater (T. cupido) tail-shakes prior to vocalizing and tail-fans at the end of the third syllable in its booming vocalizations. In the Lesser (T. pallidicinctus) tail-fanning occurs during the first syllable of the vocalization. A hybrid tail-fanned during the first (as in Lesser) and third (as in Greater) syllables of the vocalizations. The hybrid did not tail-shake and was thus similar to the Lesser.

Duration of the displays in the Greater and Lesser Prairie Chickens was 2.8 versus 0.6 seconds respectively. The hybrid’s display directed at a female (= Epigamic display) lasted 1.22 ± 0.24 seconds. When performed solitarily (= Antaposematic display), the display lasted 1.11 ± 0.29 seconds. Thus directed and undirected displays were intermediate in duration between the two parental types.

Doves
Lade and Thorpe (1964) studied the coos of five species of doves and their hybrids. Species crossed included the Barbary or Ring Dove (Streptopelia risoria), Turtle Dove (S. turtur), Collared Dove (S. decaocto), Necklace Dove (S. chinensisT and Senegal Dove (S. senegalensis).

Cross-fostering experiments failed to change the coos in any way, indicating that dove calls are genetically coded. In the Ring Dove, vocal signals developed normally even in individuals deafened soon after hatching (Konishi and Nottebohm 1969).

Lade and Thorpe (1964) concluded that specific rhythms of the
songs (coos) of doves were coded in the central nervous system. The effect of hybridization on the rhythm varied. In some crosses (e.g. Senegal x Barbary) a complete breakdown of the coding ensued so that only simple monosyllabic coos were produced by the F11 hybrids. Other hybrids (e.g. Barbary x Collared) produced coos which were rhythmically intermediate or resembled one parent. In back-crossed F2 hybrids, disorganization of the rhythm seemed to have disappeared, and the dominant species rhythm appeared in almost normal form.

Whitman (1919) studied the intervals between perch-coos of Zenaidura macroura, Streptopelia risoria and their hybrids. He noted that fifteen seconds usually elapsed between each perch-coo of Zenaidura. A Zenaidura x Streptopelia hybrid called 17 times in 20 seconds, or even once per second, a rhythm similar to the Streptopelia parent.

Carduelid Finches
Syllablic structures in the songs of canaries and Greenfinches are learned. However, Guttinger (1979) has found that rhythmic structure (temporal organization) in Greenfinch song is innate.

Syllables of both Carduelid species are grouped together into units he calls tours. In canaries, groups of tours may be separated into phrases by intervals exceeding 1 – 2 seconds of silence. Thus a histogram of intervals between tours in canary song is bimodal with a break at 1 second. Songs of Greenfinches are not broken up into phrases, and silent intervals between tours are modal in distribution. A Greenfinch raised by canaries learned canary syllables but sang them in a Greenfinch tempo, i.e. with the absence of phrases. A hybrid Greenfinch x Goldfinch likewise learned canary syllables, but temporal organization was Greenfinch-like, i.e. silent units were modal (Guttinger et al, 1978). However, whereas intervals between tours shorter than 0.5 seconds are rare in Greenfinch song, they are frequent in songs of canaries and the hybrid Greenfinch x Goldfinch.

Estrildid Finches
Cross-fostering experiments have shown that social calls of Grass Finches (Poephilae) and Mannikins (Lonchurae) develop normally independently of learning (Guttinger and Nicolai 1973, Zann 1975). Nestlings of the African Mannikins of the Spermestes/Odontospiza group are distinguished from the Asiatic Lonchura on the basis of begging calls. In the former, begging calls are bisyllabic, whereas in the latter they are monosyllabic. Guttinger’s (1970) spectrograms of begging calls of a Spermestes bicolor x Lonchura striata hybrid indicate that they are bisyllabic as in Spermestes.

Harrison (1962) described the contact call of the Bengalese Finch (L. striata) as a dry “tritt”, and that of the African Silver-bill (Euodice cantans) as a thin-shrill “psit”. The call of a hybrid E. cantans x L. striata was described as a nasal, low-pitched “kent”, thus resembling neither parent. Baptista (personal observation) has obtained spectrographic evidence to support Harrison’s description. Besides being lower pitched, contact calls of the hybrid are harmonically rich whereas those of the parental forms are relatively pure-toned, accounting for the audible differences in tonal quality.

Hummingbirds
A number of authors have suggested that the loud sounds produced at the culmination of the nuptial (aerial) dive displays of Calypte hummingbirds are produced by the specialized outermost rectrices. However, recent spectrographic evidence indicates that these sounds are mostly, if not entirely, vocal (Wells et al, 1978, Baptista and Matsin 1980). The dive sound of the Anna (C. Anna) hummer are harmonically rich, those of the Costa (C. Costae) and their hybrid are pure toned. Dive sounds of the Costa are longer in duration and higher pitched than those of Anna. Dive sounds of their hybrid are intermediate in pitch and duration between the two species.

Crowing in Phasianids
Stadie (1967) analyzed the crows of Ring-neck Pheasant x domestic cock hybrids. The crow of Gallus consisted of four indistinct syllables, with the fourth somewhat lengthened and sometimes emphasized. The crow of Phasianus consisted of two distinct “hoarse-sounding” syllables. Stadie found that not all hybrids crowed, and those that crowed rarely did so. As in Phasianus, crows of hybrids were confined to the spring. Crows of hybrids consisted of four syllables as in Gallus, but as in Phasianus syllables were clear-cut and no syllable was emphasized.

Crows of Gallus sonneratii consisted of ±5 syllables (rarely four). There was a pause between the second and third syllable, and the second and fourth syllables were emphasized. Crows of hybrids between G. sonneratii x G. gallus consisted of 3 to 6 syllables. In most cases syllables were monotoned and a little hoarse-sounding and thus unlike either parent. All hybrids sometimes had a long introductory call preceding crowing found in sonneratii but not in gallus.

Courtship

Ducks

Ethologically, ducks (Anatidae) have been the best studied of any avian group (review in Johnsgard 1965). The early studies on the behavior of duck hybrids by Lorenz (1941, 1971), Ramsay (1961) and von de Wall (1963) have been followed by two important studies: namely, those of Sharpe and Johnsgard (1966) and Kaltenhauser (1971).

Kaltenhauser (1971) studied the behavior of fifteen Anas species and their hybrids. These species possess a number of display postures in common (see, e.g. Figure 2) with some species specific variations. Some postures are absent or rare in some species (see discussion in Lorenz 1967:92 on rare behavior in ducks). Some are obligately coupled motor sequences. For example, the male Mandarin Duck (Aix galericulata) has a ritual drinking display which is always followed by sham-preening. During sham-preening he touches the raised wing “flag” on the side immediately facing the female (Lorenz 1971). Kaltenhauser (1971) chose four postures which exhibited “typical intensity” for analysis: grunt-whistle, headup-tailup, downup, and bridling. (Figure 2).

Kaltenhauser found that when two species had a display posture in common, the hybrid would exhibit the posture in an intermediate form. Sometimes a behavior was rare in one species or was present only during behavioral ontogeny, or during contexts other than courtship. Hybrids involving one such parent would exhibit the posture in an intermediate form, but during courtship as in one parent. For example, bridling is performed during courtship and as a postcopulatory display in A. flavirostris. In A. platyrhynchos, bridling is performed only by juneviles during behavioral ontogeny, or as a postcopulatory display in adults. Hybrids between these two species performed an intermediate form of bridling both as courtship and as a postcopulatory display.

Sometimes behaviors that were rare or absent in the parental forms were present in the hybrids. For example, A. acuta x A. spinicauda hybrids performed downup and bridling displays both of which were apparently absent in the parents. A. platyrhyncho x A. strepera backcrossed to A. platyrhynchos performed bridling displays absent in the parental forms as well as Fl hybrids.

The inheritance of coupled sequences was also studied. She found that coupled sequences found in both parents appeared also in Fl hybrids, albeit sometimes rarely (e.g. the grunt-whistle coupled with the headup-tailup in A. acuta x A. castanea, Table 1). If a coupled sequence was absent in one parent, it was also absent in the hybrids (e.g. A. penelope x A. strepera and A. castanea x A. spinicauda). The time intervals between coupled sequences were often longer in hybrids than in their parents. For example, the interval between grunt-whistle – headup-tailup sequences in A. crecca and A. flavirostris ranged from 0.8 – 1.8 sec. (x = 1.3 seconds); in their hybrids, intervals ranged from 1.6 to 2.0 seconds (x-bar = 1.8 seconds). In two cases hybrids showed coupled sequences not found in either parent. Indeed, in A. sibilatrix x A. strepera, two new coupled sequences appeared (Table 1).

Sharpe and Johnsgard (1966) (see also Johnsgard 1967) studied the behavior of hybrids between A. platyrhynchos and A. acuta and correlated these with plumage inheritance. Both mallard and pintail perform grunt-whistle and headup-tailup. However, only the mallard performs the downup, whereas only the pintail performs the burp (Figure 2). Nod-swim is commonly performed by the mallard but is rudimentary in the pintail. In Fl hybrids the downup is lacking and burp and nod-swim appear in reduced form. F2’s exhibited plumage and behavioral variation ranging from almost pure pintail to almost pure mallard-like. In the most mallard-like F2’s, downup and nod-swim are well developed. The most pintail-like F2’s performed well-developed burp displays. Hybrid scores were constructed for plumage and behavior, and a positive correlation was found between inheritance of plumage and behavioral characteristics. The considerable segregation and independent assortment observed suggested to the authors that both plumage and behavior were under fairly simple genetic control, probably involving relatively few genes.

No abnormal coupled sequences were found in F2’s. However, an F3 linked a bridling (postcopulatory) display with a headup-tailup social display.
Lind and Poulson (1963) described the courtship displays of a Goosander x Shelduck hybrid. They found that during the ritualized drinking display, Goosanders normally held their head up 90 degrees with the horizontal, Shelducks held their heads at 45 degree, and the hybrid held its at an angle intermediate between the two parents.

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Doves
Davies (1970) studied bowing displays in Streptopelia doves and their hybrids. These were the same five species studied by Lade and Thorpe (1964) who described their vocalizations. Bowing displays were performed only by males, were directed at either sex, were species specific, and involved an alternate upright and crouched posture which exhibited “typical intensity”. Bowing displays of hybrids also exhibited typical intensity. Some of the components

of the bows were intermediate between the parents, some resembled one parent closely, and others exceeded either parent. For example, the display of Collared x Barbary Dove hybrids (Figure 3) resembled the Barbary parent at the top of the bow in that the head was not held up as high as the Collared parent. However, the hybrid bowed lower than either parent. The exaggerated movements were lost in the F2 generation.
Similar results were obtained when Davies examined frequency of bouts of bowing, e.g. Turtle x Barbary hybrids bowed at a frequency intermediate between the parents, but frequency of bowing in Necklace x Barbary hybrids approximated the Barbary parent.

Hummingbirds
North American Hummingbirds of the genus Calypte exhibit two displays during territoriality or courtship: (1) a static display during which individuals sing from a perch and (2) a dynamic display which involves singing and species specific movements through the air. Selasphorus Hummingbirds only perform dynamic displays. On the Palos Verdes Peninsula of Southern California, three species of hummingbirds, the Anna (Calypte anna), the Costa (C. costae) and the Allen (Selasphorus sasin) breed sympatrically and overlap extensively in breeding season. Occasional hybrids have been found and their displays analyzed (Wells, Bradley and Baptista 1978, Wells and Baptista 1979).

In the dynamic display of the Anna Hummingbird, the male sings while hovering some 25 to 50 meters above the female. He then ascends, continually looking down (bill pointed down). He pauses at the top of the climb and may sing once more, and then dives, swooping out of his dive just over the female and produces a loud chirp sound. In contrast, male Costa Hummingbirds climb up into the air over the female with bill pointed up, never sing, never pause at the top, then dive and produce a loud whistling sound. One male Costa Hummer was observed tracing a large horizontal circle at the top of his climb just prior to diving. Hybrid Anna x Costa hybrids perform the aerial display with the bill pointed down like the Anna parent, climb up into the air in silence like the Costa, then trace a large horizontal circle in the air before diving and making a sharp but short peek sound. These hybrids also possess a static display wherein they sing songs identical to the Anna parent. It is conceivable, however, that songs are learned in hummingbirds (see discussion in Mirsky, 1976).

Allen Hummingbirds begin their aerial display by flying back and forth tracing a pendulum path over the female. At each end of the pendulum, a high-pitched cricket-like chirping is produced at between 7.5 and 12 kHz. After a number of these horizontal arcs, the hummer climbs in silence 75 to 100 feet into the air, describing spirals or undulations all the way to the top. He never pauses at the top but dives immediately. As he breaks his dive over the female, he produces a loud sound which begins with a number of short pip sounds followed by a long whistle. A hybrid Anna x Allen climbed into the air with bill pointed down like the Anna parent, but traced an undulatory path to the top like the Allen. He sometimes paused at the top like the Anna but was silent like the Allen. The hybrid then dived and produced a loud peek sound at the bottom of the dive. The dive sound was spectrographically like the Allen in harmonic structure, but short in duration like the Anna parent. The pendulum display of the Allen Hummingbird was absent in the hybrid. The hybrid also had a static display like its Anna parent, but its song was unlike any hummingbird known.

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Figure 3.
Bowing displays of doves:
a. Collared Dove
b. Barbary Dove
c. Hybrid Collared x Barbary Dove
After Davis, 1970.

Estrildid Finches
Eisner (1958) described the displays of Bengalese Finch (Lonchura striata) x African Silverbill (Euodice cantans) hybrids, and Harrison (1962) described displays of the reciprocal cross. I made observations additional to those of Harrison (1962) on Silverbill x Bengalese crosses, and extended their observation to three other crosses involving Estrildid Finches (Table 2).

Males of both Silverbill species (E. cantans and E. malabarica) perform a precopulatory display holding a grass stem by one end, all of the body feathers sleeked, tail shut and pointed down, and inverted curtsies (a bobbing movement in which the main component is an upward thrust imparted by a straightening of the legs). Sometimes birds perform ritualized nest-building movements while holding the straw as if molding an invisible nest roof (= U-shaped movements). Hybrids from three crosses involving Silverbills as one parent, performed all the components of the straw display (Table 2), but with some of the feather postures of the other parent. For example, when performed at high intensity, Silverbill X Bengalese crosses spread their tail and fluff their belly feathers like the Bengalese parent, but tails are always pointed down like the Silverbill parent. Silverbill x Zebra Finch (Poephila guttata) crosses always ruffle their nape and belly feathers like the Zebra parent. Some Spice Finches (L. punctulata) sometimes perform inverted curtsies during their precopulatory display as do Spice Finch x Bengalese hybrids. Spice and Bengalese Finches bow before singing at strangers or prior to precopulatory displays. When isolated and then presented with a female, two hybrid combinations involving Spice and Bengalese Finches also bow prior to singing.

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Figure 4.
Peering behavior in Lonchura punctulata. The bird at right is peering at the singing male at left. After Morris, 1958.

Some Estrildid Finches also perform a display known as peering when one bird sings and other members of the social flock sleek their feathers and appear to be listening intently at close range (Figure 4). This display may serve to strengthen flock bonds (Immelmann 1965). All crosses involving peering (E. malabarica,
E. cantans, L. punctulata) and non-peering (P. guttata, L. striata) species performed the peering display (Table 2).

Viduine Finches
Payne (1980) studied displays of Village Indigobird (Vidua chalybeata) x Paradise Whydah (V. paradisaea) hybrids in Lonchinvar Park, Zambia. Viduine Finches usually oviposit in nests of specific host Estrildid Finches. The parasite then learns the songs of its host, and later learns the song of its own species. Since the hybrids sang songs of Paradise Whydahs and Melba Finches (their host), Payne suggested that the crosses he studied must have involved the Paradise Whydah as the female.
Paradise Whydahs perform an elaborate aerial display which was absent in Indigo birds and their hybrids. One observation was made of a hybrid performing a courtship display termed a “fast sideways head bent”, given by Paradise Whydahs. Indig obirds seem to sing and display throughout the day, whereas Whydahs and the hybrids sang only during mid-morning and mid-afternoon. Hybrids were also similar to Indigo-birds in that most songs were delivered from one tree rather than from several singing trees as in Whydahs.

Absence of Courtship Displays
A number of authors have reported the absence of displays in some intergeneric hybrids between distantly related species. Asmundson and Lorenz (1955) produced Ring-neck Pheasant x turkey hybrids by artificial insemination. They reported the absence of eggs or semen in the hybrids and never observed courtship. Huxley (1941) kept an impeyan x Lady Amherst Pheasant hybrid with a female Impeyan and reported no trace of sexual interest in the hybrid. Poulson (1950) studied a domestic duck x Greylag Goose hybrid. Despite injections of testosterone, no courtship displays were observed. Poulson interpreted this as an absence of response from the nervous system.

Nest Building in Agapornis Lovebird Hybrids
Nest building behavior in the African Peachface (Agapornis roseiocollis) and Fisher’s (A. [personata] fischeri) Lovebirds (Psittacidae) and their hybrids was studied by Dilger (1962a, 1962b) and his student Buckley (1969). A. fischeri cuts strips of paper about 7 inches long and 3/8 inches wide and carries these with the bill one at a time into the nest box. Sticks, strips of bark or leaves are also utilized in their nest. A. roseicollis cuts strips 3 to 4 inches long, then tucks several pieces under the feathers of the rump and lower back and thus carries them into the nest box (Figure 5).

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Figure 5.
Nest building by Agapornis roseicollis. The female is tucking a cut strip of paper under its rump feathers. After Dilger 1962a.

Hybrids between roseicollis and fischeri cut paper strips that were longer than those of either parent. Strip-cutting is normally practiced by females of both species: occasional males of roseicollis cut strips, and males of fischeri almost never thus perform. Fl hybrid males cut and carry with greater frequency than males of either parental form. Indeed, these males were continually interfering with strip-cutting attempts of the hybrid females; males of the parental forms rarely do.
Hybrids also exhibited various conflict behavior patterns and were rarely successful in completing a tucking sequence. They might perform the proper movements of tucking, but appeared unable to let go of the strips. They might perform frequent intention movements to tuck without completing the act. Sometimes they attempted to reach the rump with the bill by running backwards. Hybrids often grasped the strips other than by either end, thus making tucking difficult. ‘They were often distracted, so that tucking attempts often passed into drinking or displacement activities (e.g. head-scratching). Sometimes inappropriate items, e.g. twigs, were tucked. This behavior, Buckley (1969) pointed out, was understandable since fischeri often incorporated twigs in their nests. Hybrids would sometimes tuck in inappropriate places, e.g. the breast. Examination of movie footage indicated that hybrids lacked the proper act sequence for successful tucking. That is, items were not correctly placed in the rump. indeed, an important component called “tremble-shoving” was lacking in the hybrids.

Dilger (1962a) observed that hybrids would gradually abandon the abortive tucking attempts in favor of carrying in the bill. After three years, tucking became very infrequent and behavior was very much like A. fischeri. However, Buckley (1969) studied a mixed batch of hybrids ranging from six months to four years old and reported that they could not be reliably divided into age groups on the basis of relative tucking and bill-carrying frequencies. There appears, thus, to be individual variation in hybrids in their abilities to learn to bill-carry. All Buckley’s birds showed both rump-tucking and bill-carrying behavior, with bill-carrying as the last resort.

DISCUSSION
In this paper I have brought together some of the literature on the behavior of domesticated birds, showing how man has modified some of these behavior patterns including social behavior, displays and wildness. Some of these behavior patterns are inherited in crosses between strains in the same way as in Fl hybrids between species. Some of these behavior patterns are clearly mal-adaptive and are permitted to survive only because of protection by man. For example, the aberrant flight behavior of the Swing Pouter Pigeon causing him to lose altitude and the clapping behavior which causes premature fraying of the primaries would probably have been selected out long ago by predators had these mutations occurred in the wild (Nicolai 1976). Since courtship behavior patterns may function as species isolating mechanisms, extreme modifications of these behaviors may present an individual from acquiring a mate in the wild, and thus also be eliminated from natural populations.

In the second portion of this paper I have summarized studies on the behavior of artificial and natural avian hybrids involving five different orders: Galliformes, Anseriformes, Columbiformes, Apodiformes and Passeriformes. Since most Fl hybrids are sterile, .few of these studies were carried on to the F2 generation. The studies by Sharpe and Johnsgard (1966) on duck hybrids and Entrikin and Erway (1972) on rolling in domestic pigeon breeds are especially noteworthy, both because of the excellent analyses and sample sizes involved.

Frank (1974) and Payne (1980) have summarized the various trends in the behavior of hybrids which are as follows:
(1) Hybrids may acquire behavior patterns similar to one or both of the parental forms (e.g. foraging behavior patterns) suggesting genetic dominance.
(2) Hybrids may perform behavior patterns intermediate between the two parental forms (e.g. duration and pitch of distress calls of chicken x pheasant hybrids).
(3) Hybrids may perform exaggerated versions of the parental behavior patterns (e.g. amplitude of bow in the Collared x Barbary Dove hybrids).
(4) Behavior patterns rare in the parental forms may be common in the hybrid (e.g. horizontal circle flight of the Anna x Costa Hummingbird hybrid).
(5) Behavior of hybrids may be incomplete (intention movements), e.g. tucking attempts of Agapornis hybrids, or may be otherwise disorganized (e.g. cooing of S. senegalensis x S. risoria hybrids) due to alteration of the genome.
(6) Complex behavior sequences may be reorganized or new combinations of behavior components may arise (e.g. movements and feather-postures of Estrildids are recombined in the hybrids), suggesting segregation and recombination.
(7) Behavior patterns may appear in the hybrid not found in either parent, e.g. Agapornis hybrids tucking in the breast rather than the rump.
(8) Behavior patterns of both parents may fail to appear in the hybrid (e.g. absence of displays in some hybrids). Hinde (1956) has suggested that this is due to the general developmental breakdown of the hybrid.

A number of authors have interpreted some of the behavior of hybrids as a recapitulation of ancestral behavior patterns (Lind and Poulson 1963, Dilger 1962b, Buckley 1969). For example, Dilger and Buckley noted that some primitive Agapornis species as well as the related Blue-crowned Hanging Parrot (Loriculus galgulus) regularly tuck nest material in the breast. Breast-tucking in Agapornis hybrids, found in neither parental form, has been interpreted as an atavistic behavior which has emerged as a result of changes in the genome. Franck (1974) has taken exception to this idea suggesting that the altered genotype has perhaps disrupted the parental behavior pattern resulting in a behavior accidentally resembling the primitive forms.

Rare parental behavior occurring commonly in hybrids have been interpreted as a lowering of thresholds to perform these behavior patterns due to the altered genome (Kaltenhauser 1971). Indeed, Manning (1963) has suggested that many evolutionary changes in behavior are due to altered thresholds to perform these behaviors. Franck (1974) argues against this, pointing out that we have as yet no actual understanding of the action of genes on behavior or a physiological understanding of the changes that can be described as threshold changes.

SUMMARY
Studies on the behavior of domesticated birds were reviewed. During the process of domestication man has selected for certain behavioral, traits including tameness, changes in social behavior, qualities of voice and changes in courtship patterns. Since the ancestral wild forms still exist, comparison of these behaviors in wild and domesticated forms give us some ideas as to how natural selection operates to change dispositions to perform behavior patterns. Certain behavior patterns are inherited in crosses between domestic strains in much the same way as inheritance of behavior in Fl interspecific hybrids.

Studies on the behavior of natural and artificial (laboratory produced) interspecific hybrids were surveyed. Heritable characteristics included maintenance activity patterns, quality of voice, courtship patterns, and nest-building behavior. Since Fl hybrids are usually sterile, few studies continued on to F2 generations. Behavior patterns in hybrids may (1) resemble one parent, (2) be present in intermediate form, (3) resemble neither parent or (4) be absent altogether. Behavior rare in parental forms but common in hybrids have been interpreted as changed thresholds to perform these behavior patterns or recapitulation of ancestral behavioral traits due to altered genomes. Alternate interpretations of these data are discussed.

ACKNOWLEDGMENTS
I thank Kerry Muller and Robert B. Payne for their critical comments on the manuscript. Kerry Muller also called my attention to some literature on behavior of domestic pigeons. Roland Sossinka assisted me with the translation of some of the German literature.

REFERENCES
Asmundson, V.S. and F.W. Lorenz. 1955. Pheasant-turkey hybrids. Science, 121:307-308.
Baptista, L.F. 1973a. Song mimesis by a captive Gouldian Finch. Auk. 90:891-894.
1973b. On courtship displays and the taxonomic position of the Grey-headed Silverbill. Avicult. Mag. 79: 148-154.
Baptista, L.F. and M. Abs. 1981. Vocalizations of the Rock Dove and some Domestic Pigeon breeds. In M. Abs (ed.) Behavior and Physiology of the Pigeon. Academic Press. In press.
Baptista, L.F. and M. Matsui. 1979. The source of the dive-noise of the Anna Hummingbird. Condor, 81:87-89.
Buckley, P.A. 1969. Disruption of species – typical behavior patterns
in Fl hybrid Agapornis parrots. Z. Tierpsychol. 26:737-743.
Clark, G.A., Jr. 1973. Holding food with the feet in passerines. Bird-banding, 44:91-99.
Crawford, J.A. 1978. Morphology and behavior of Greater x Lesser Prairie Chicken Hybrids. The Southwestern Nat., 23:591-596.
Davies, S.J.J.F. 1970. Patterns of inheritance in the bowing display and associated behaviour of some hybrid Streptopelia Doves. Behaviour, 36:187-214.
Darwin, C. 1875. The variation of animals and plants under domestication. 2nd ed. London, J. Murray.
Desforges, M.F. and D.G.M. Wood-Gush. 1975a. A behavioral comparison of domestic and Mallard Ducks: habituation and flight reactions. Anim. Behay., 23:692-697.
3 1975b. A behavioral comparison of domestic and Mallard Ducks: spatial relationships in small flocks. Anim. Behay., 23:698-705.
1976. Behavioral comparison of Aylesbury and Mallard Ducks: sexual behaviour. Anim. Behay., 24:391-397.
Dilger, W.C. 1962a. The behavior of lovebirds. Sci. Amer., 206: 88-98.
1962b. Behavior and genetics. In E. L. Bliss (ed.)
Roots of behavior. Harper Bros., N.Y.
5
Edrich, W. and W.T. Keeton, 1977. A comparison of homing behavior
in feral and homing pigeons. Z. Tierpsychol., 44:389-401.
Eisner, E. 1958. Bengalese Finch x Silverbill hybrids. Avicult. Mag., 64:5-1-54,
Entrikin, R.K. and L.C. Erway. 1972. A genetic Investigation of Roller and Tumbler Pigeons. Journal of Heredity, 63:351-354.
Fennel, R.A. 1945. The relation between heredity, sexual activity and training to dominance – subordination in game cocks. Am. Nat., 79:142-151 .
Franck, D. 1974. The genetic basis of evolutionary changes in behavior patterns. In J.H.F. Van Abeleen (ed.) The genetics of behavior. pp 119-140. Amsterdam, Holland.
Goodwin, D. 1965. Instructions to young ornithologists IV. Domestic Birds. London Museum Press, Ltd.
Gould, J.L. 1974. Genetics and molecular ethology. Z. Tierpsychol . , 36:267-292.
Guttinqer, H.R. 1970. Zur Evolution von Verhaltenswiersen und Lautausserungen bei Prachtfinken (Estrildidae). Z. Tierpsychol., 27:1011-1075.
1979. The Integration of Learnt and Genetically
Programmed Behaviour: A study of Hierarchical Organization in Songs of Canaries, Greenfinches and their hybrids. Z. Tierpsychol., 49:2811-31-303.
Guttinger, H.R. and J. Nicolai. 1973. Struktur und funktion der
rufe bei prachtfinken (Estrildidae). Tierpsychol., 33:319-334.
J. Wolffgramm and T. Thimm. 1978. The relationship between species specific song programs and individual learning in song birds. Behaviour, 65:241-262,
Harrison, C.J.O. 1962, A Silverbill x Bengalese Finch hybrid. Aviculture Mag., 68:30-33.
Hinde, R.A. 1956. The behaviour of certain Cardueline Fl interspecies hybrids. Behaviour, 9:202-213.
Howes, J.R. 1964. Japanese Quail as found in Japan. The Quail Quarterly, 1:21-30.
Huxley, J.S. 1941. Genetic interactions in a hybrid pheasant. Zool. Soc. (London) Proc. IIIA:41-43.
Immelmann, K. 1965, Australian Finches. Angus and Robertson. Melbourne.
1969. Song development in the Zebra Finch and other estrildid finches. In R. A. Hinde (ed.) Bird Vocalizations. Cambridge, England. pp 61-74.
1976. Einfuhrung in die Verhaltensforschung. Verlag
Paul Parey. Berlin.
, A. Piltz, and R. Sossinka. 1977. Experimentelle untersuchungen zur Bedeutung der Rachenzeichnung junger Zebrafinken. Z. Tierpsychol., 45:210-218.
Johnsgard, P.A. 1965. Handbook of Waterfowl Behavior. Cornell University Press.
1967. Animal Behavior. W. C. Brown Co., Iowa,
Kaltenhauser, D. 1971. Uber Evolutionsvorgange in der Schwimmentenbalz. Z. Tierpsychol., 29:481-540,
Konishi, M. 1963. The role of auditory feedback in the vocal behaviour of the domestic fowl. Z. Tierpsychol., 20:349-367.
and F. Nottebohm. 1969. Experimental studies in the ontogeny
of avian vocalizations. In R. A. Hinde (ed.) Bird Vocalizations, Cambridge Univ. Press. pp 29-48.
Kovach, J.K. 1974, The Behaviour of Japanese Quail: Review of Literature from a Bio-ethological Perspective. Applied Animal Ethology, 1:77-102.
Kruijt, J.P. 1964. Ontogeny of social behaviour in Burmese Red Junglefowl, Behaviour, Supplement 12.
Lade, Barbara I. and W.H. Thorpe. 1964. Dove songs as innately coded patterns of specific behaviour. Nature, 202:366-368.
Leopold, A.S. 1944. The nature of heritable wildness in turkeys. Condor, 46:133-197.
Levi, W.M. 1965. Encyclopedia of Pigeon Breeds. T.F.H. Publications, Inc., N.J.
Lind, H. and H. Poulson, 1963. On the morphology and behaviour of a hybrid between Goosander and Shelduck (Mergus merganser L. x Tadorna tadorna). Z. Tierpsychol., 20:558-569.
Lorenz, K. 1941. Vergleichende Bewegungsstudien an Anatinen. J. Ornithol., 89 (Suppl.):194-294.
1969. Evolution and modification of behavior. Univ.
of Chicago Press. i 1971. Studies in Animal and Human Behavior II. Harvar
University Press, Cambridge, Mass.
Manning, A. 1963. Evolutionary changes and behavior genetics. Proc.
XIth Internat. Cong. Genetics, 3 (Oxford, Pergamon):807-815.
1972. An Introduction to Animal Behavior, 2nd Edition.
E. Arnold Ltd., London
Marler, P. 1959. Developments in the study of animal communication. Ch. 4 In P. R. Bell (ed.) Darwin’s Biological Work: Some Aspects Reconsidered. Cambridge Univ. Press, England.
, and M. S. Waser. 1977. Role of auditory feedback in canary song development. J. Comp. Physic. Psychl., 91:8-16,
McGrath, T.A., M.D. Shalter, and W.M. Schleidt, 1972. Analysis of distress calls of chicken x pheasant hybrids. Nature,, 237:47-48,
Mirsky, E.N. 1976. Song divergence in hummingbird and junco populations on Guadalupe Island, Condor, 78:230-235.
Moreau, R.E., and P. Wayre. 1968. On the Palearctic Quails. Ardea, 56:209-227.
Morris, D. 1958. The comparative ethology of Grassfinches (Erythrurae) and Mannikins (Amadinae). Proc. Zool. Soc. London, 131:389-439.
Mundinger, P. 1970. Vocal imitation and individual recognition of finch calls. Science 168: 480-482.
Nicolai, J. 1960. Verhaltensstudien an einigen afrikanischen und palaarktischen Girlitzen. Zool. jb., 87:317-362.
1964. Der Brutparasitismus der Viduinae als ethologisches problem. Z. Tierpsychol., 21:129-204.
1967. Vogelhaltung – Vogelpflege. Kosmos,
Stuttgart, W. Germany,
1976. Evolutive Neuerungen in der Balz von Haustautienrassen. (Columba livia var. domestica) als Ergebnis menschlicher Zuchtwahl. Z. Tierpsychol., 40:225-243.
Payne, R.B. 1980. Behavior and songs of hybrid parasitic finches. Auk. 97:118-134.
Poulson, H. 1950. Morphological and ethological notes on a hybrid between a domestic duck and a domestic goose. Behaviour., 3:99-104.
1959. Song learning in the domestic canary. Z. f. Tierpsychol., 16:1173-178.
Ramsay, A.C. 1961. Behaviour of some hybrids in the mallard group. Anim. Behay., 9:104.
Sharpe, R.S., and P.A. Johnsgard. 1966. Inheritance of behavioral characters in F2 Mallard x Pintail (Anas platyrhynchos L. x Anas acuta L.) hybrids. Behaviour, 27:259-272.
Siegel, P.B. 1965. Genetics of behavior: selection for mating ability in chickens. Genetics, 52:1269-1277.
, R. E. Phillips and E. F. Folsom. 1964. Genetic variation in the crow of adult chickens. Behaviour, 24:229-235.
Stadie, C. 1967. Verhaltensweisen von Gattungsbastarden Phasianus colchinus x Gallus gallus F. domestics im Vergleich mit deren der Ausgangarten. Zool. Anz., 31st Suppl.
1969. Vergleichende Beobachtungen an Verhaltensweisen
verschiedener Wildhuhnarten der Gattung Gallus. Zool. Anz., 183:13-30.
Von de Wall, W. 1963. Bewegungsstudien an anatinen. J. Ornithol. 104:1-115.
Waser, M.S. and P. Marley. 1977. Song learning in canaries. J. Comp. Physiol. Psych., 91:1-7.
, R. A. Bradley and L.F. Baptista. 1978. Hybridization
in Calypte hummingbirds. Auk, 95:537-549.
Wells, S. and L.F. Baptista. 1979. Displays of an Anna x Allen Hummingbird hybrid. Wilson Bulletin, 91:524-532.
Whitman, C.O. 1919. The behavior of pigeons. Posthumous Works, Vol. III, Carnegie Institute, Washington, D.C.
Wood-Gush, D.G.M. 1960. A study of sex drive of two strains of cockerels through three generations. Anim. Behay., 8:43-53.
Yamashina, Y. 1961. Quail breeding in Japan. J. Bombay Nat. Hist. Soc. 58:216-22.
Zann, R. 1975. Inter- and intraspecific variation in the calls of the three species of Grassfinches of the subgenus Poephila (Gould) (Estrildidae). Z. Tierpsychol., 39:85-125.

ROSELLAS

By Gene Hall
Fortune Glen Aviaries

IFCB 1978

I have been involved for the past twelve years, 1966 through 1978, with the genus commonly known as Rosellas (Platycercus). Indigenous to Australia, they comprise a group of eight species which are quite colorful, not excessively noisy or destructive and fairly willing to reproduce in captive situations. They range from ten to fourteen inches long, medium in body size, and are of the typical parakeet type. They are fairly active, capable of rapid flight and move easily on the floor of the aviaries while feeding or bathing.

All of the species have cheek patches and reasonably long full tails. Hence, the descriptive term “Broadtails” and the feathers of the mantle, back and wings are black with a bordering of another color which produces a scalloped effect. An attempt to describe each species would be lengthy and laborious as all of them are complicated in their coloring. There are many excellent books available containing photographs of them, or better yet, a visit to a collection will be worth a thousand words.

Five of the species come from the eastern portion of Australia, one from Tasmania, one from the northern area and one from the southwestern area. Both sexes are colored alike in all species except for the Stanleys. In some cases very slight differences in color intensity does exist between sexes.

Sexing can be quite difficult unless you are fortunate enough to deal with specimens that show distinctive physical differences. Ideally, the size and shape of the head will often offer clues. The males should have a larger head with plenty of brow or frontal while the hen’s will be lacking in brow or frontal. Nevertheless, a side view study of both is helpful. H. D. Groen in his book “Australian Parakeets” does an excellent job indicating differences photographically. When viewed from the front, the mandible of the male should be larger and broader than that of the hen. As they mature, a close study of their behavior will be most informative with the males quite aggressive with each other. I have not found the wing stripe to be accurate enough to depend on in determining sex. The Stanley shows a definite difference in color with the hen lacking much of the red seen on the male and the yellow cheek patch is smaller and less colorful.

In an effort to understand the needs of Rosellas, I have read all of the publications I could find not only concerning the birds and their behavioral habits but any information regarding their native habitats. With the quarantine regulations permitting their entry in 1968 I was able to obtain good European stock to enlarge the breeding program. Our move to a rural San Diego location from Los Angeles area in 1972 gave us more room to expand, cleaner air, untreated well water, natural open surroundings and, most important, climate like that of their native habitat.

We have bred four species, the Eastern Platycerus eximus; blue, Platycerus adscitus; Crimson, Platycerus elegans; and Stanley, Platycerus icterotis; and recently acquired the Adelaide, Platycerus adelaidae. The crimson or pennant has been our main interest from the beginning and has produced approximately eighty youngsters at the present location.

Because of the area’s temperatures, average lows of 25 and highs of 110, the aviaries are designed without closed shelters or heat. The aviaries are four feet wide and 125 feet long, running north to south and divided into six units ranging from 16 to 25 feet per unit. The shelter end of each unit backs to the north, blocking the cold winter winds that blow down the valley and thus becomes a solid end to the flight behind. The closed sides of the shelter extend 4 feet on the side, allowing a 4 x 4 foot area for nest boxes, feeding arrangements and privacy. The sides of the flight area are 112 x 112 inch welded wire which allows plenty of sun both early and late in the day. During the summer the prevailing winds from midday on are from the ocean ten miles to the west and keep the temperatures quite comfortable. The roof is fully covered, providing good protection from sun and rain but, just as important, limits the visibility of the many varieties of birds of prey in our valley.

Perches of eucalyptus branches are placed on each end of the aviaries, allowing the maximum unobstructed flight, thereby maintaining strength, vigor and fertility. This also allows them to satisfy their emotional need for early and late flights as they might in the wild in their quest for feed and water. The crimsons are kept in the 25 foot flights, and the rest in sixteens. This end on end arrangement of aviaries has eliminated the problems previously experienced and added no new ones except construction costs which has been absorbed by increased productivity. Fighting between males in adjoining aviaries over territorial rights as well as the courting of the other’s hen has been eliminated. Therefore, the pairs must concern themselves with each other without distraction.

As the breeding season approaches, the males begin the courting calls which stimulates the entire group which all are close enough to hear but unable to see. These long units are separated by 20 feet which are planted to limit visibility of other units and provide natural surroundings. This has resulted in a greater percentage of pairs working. The end of the flight being solid has put an end to broken necks caused by flying into wire as previously experienced in flights with wire ends.

The seeds provided are on a free choice basis: straight canary, small finch millet, large proso millet, hulled oats, safflower, sunflower, flax and niger (or thistle) and poppy (or maw). A small portion of the whole wheat bread, apple and cooked feed corn are offered daily. The seeds are all offered in dishes and feeders approximately three feet above the floor, and the bread, apple and corn fed on the floor as these birds enjoy running about the flight looking for food. Granite grit and natural cuttlebone are also placed in each flight. Regarding the feeding of greens, we feel that only home grown greens are to be considered safe. The many insecticides used today are either directly sprayed on the plants or are included in the fertilization process, and all must be considered as unsafe. Home grown swiss chard and New Zealand spinach maintain themselves for years without additional planting and the birds thrive on a small amount of these dark greens. Grated carrot is of great benefit and readily acceptable. Vitamins in some form are considered to be beneficial when added to their diet. The above is a practical diet from a nutritional point of view and is not time consuming. More elaborate diets can be fed, but unless it is possible to be consistent it is not always practical. Fresh eucalyptus growth is provided weekly and seems to provide medicinal effects both internally and externally as well as emotional satisfaction. We have not experienced external parasites since this practice was introduced. Leaves of the blue gum, Eucalyptus globulus, seem to be most satisfactory for this purpose. Entry into the flights is only necessary once a week to feed. The bread, apple and corn can be supplied without entry, and the water is changed daily from the outside. Thus, daily servicing is minimal, providing the birds with almost total privacy and freedom from stress. During the breeding season the area is closed to all but those attending to their needs on a daily basis.

The boxes furnished for nesting are of the grandfather clock type, have proved successful and are constructed from 112″ plywood, 12″ x 12″ and 24 to 48″ deep. The greater the depth, the longer the young seem to stay in the nest to mature and so are not apt to go to the floor when they fledge. The entrance hole is 3″ in diameter and 6″ from the top. A 6″ porch 2″ below the entrance hole is used rather than a perch. It allows room for the birds to sit by the entrance hole and does not become loose as a perch quite often does. As we are all aware, loose perches can be the cause of infertile eggs. The inside of the front of the nest is lined with 1/2 x 112 wire for access to the bottom of the nest. Nesting materials include a mixture of dirt, pine shavings and old eucalyptus leaves placed in the bottom of the nest to a depth of 12″ in a 48″ box. This mixture is soaked with water in late January and is ready for the hen in the middle of March. The hen will dig a depression in this mixture about 4″ to 6″ deep. She will lay every other day. Clutches usually number four to eight eggs. Incubation takes approximately eighteen to twenty-one days with the hen doing the setting. She will generally leave the nest twice daily to be fed by the male and for water. At hatching time she will leave the nest to bathe, returning to the nest quite wet, which adds moisture to the nest site. Soaking the nesting material in January is also an aid to this end. As the moisture softens the eggshell, the young are better able to escape in good time from the shell. Hens generally set fairly tight until the young are a week or more old, at which time she will leave for longer intervals. The young are fully covered with white down and cluster together. As our practice is never to disturb them at this point, it is quite often impossible to ascertain the number of young present. At approximately ten days of age the pin feathering starts and the chicks separate from this cluster.

Up to this point the male feeds the hen who, in turn, enters the nest to feed the young. By the time the chicks are two weeks old, we generally find both parents at the feed station and entering the nest to feed the young. The young will fledge at about six weeks, depending on the weather and size of clutch. It is best to leave the young with the parents for six weeks after fledging if the male will allow. This gives them a time to be thoroughly independent and the parents to imprint behavior traits. We have found that the longer the young are left with parents, the more satisfactory the results are when they are used as breeders. Our pairs have double clutched every year except 1977. The month of June, which is generally cool, was quite warm and the birds did not return to nest as in the years before.

Rosellas are capable of producing when one year of age. However, we wait until they are two years old before using them for breeding. In doing so, we have not lost hens to egg binding and our males have not been wife-beaters. They also seem to do a better job of raising their young. With the present supply of birds available today, there can be no satisfactory excuse for breeding brothers and sisters as was the practice years ago.

Acquiring stock for breeding purposes must be done selectively with the final results kept fully in mind so seldom is a pair of proven breeders sold. There are times when an aviculturist must dispose of his good pairs, but this is not a common situation. We, therefore, have acquired our share of problem birds in the past, such as the “old and sterile”, wife-beaters, egg eaters, feather pickers and poor feeders. Therefore, the only birds purchased today are in juvenile plumage from stock known to be productive.

When buying domestic-bred birds, this is a fairly simple matter. A visit to the seller not only offers the opportunity to view the number and quantity of the young produced but also a chance to see the parent stock, methods of breeding and management.

When purchasing imports, it’s a bit difficult. I would rather purchase from a large group. It indicates that the source in Europe is productive. It also gives a better choice. In either case (as well as when judging your own young), select only a small number of the group for your purposes as expanding too rapidly generally results in lowering your standards. A small group of quality specimens is far better than a larger group of lesser quality. In making your choice, we suggest picking the hens first. We place great importance on as large a hen as possible, broad in breast and shoulders, good clear eyes, calm but alert and firm in the hand.

Flighty hens are apt to break eggs and step on the young if something disturbs them while on the nest. If possible, choose hens from mothers that lay large cluthes and are successful in raising their young.

The choosing of males is a bit easier. Points of importance are: color and aggressiveness, or vigor. We try to breed size from the hen and color from the male. If the male has excellent color and is also large, so much the better.

Learn to whistle a challenge call and try it on a group of males. The males we choose will not let a challenge go unanswered.

After our choices are made, we place them in flights that are 8 x 8 feet, which allows plenty of room for exercise and growth.

Rosellas continue to grow and develop size up to twenty-four months. We hold them in these flights until they are eighteen months old. By that time, they have chosen mates and become aggressive toward each other and separation is necessary. This is a good time to re-evaluate and cull out those that have not developed to our standards. They are then ready to be moved into the breeding flights and thus will be settled in nicely by the time spring arrives. It should be pointed out that only one pair per flight is possible.

On the subject of medication, the average aviculturist is far too quick to put something in the water. Many birds are lost annually due to over-medication. Even more distressing is the use of incorrect medications in the treatment of ailing birds, which produces negative results. We agree that a well-stocked medicine chest is necessary, one that contains a complete assortment selected by your veterinarian. An excellent veterinarian in our area has become interested in our problems and has undertaken to specialize in birds. We, in turn, supply him with all literature available to us, including tape recordings of meetings and printed matter with the end result of a better program of care available to us. It is, therefore, in the best interests of a serious aviculturist to help interest and inform a veterinarian in your area so that if a real problem occurs he can take charge effectively.

Rosellas are susceptible to internal parasites and must be watched closely. The purchase of a small microscope helps in this matter. With a little practice you can determine the type of infestation and take appropriate action.

We feel that in terms of general management that all too often the aviculturists interfere with their birds. In the wild the many species have developed and prospered on their own and not until man moved in did many of their problems develop. When we place them in a captive environment we alter their lives. It is our obligation to furnish proper and adequate facilities. The birds neither read our books nor attend our meetings, and yet it is they who hold the key to the reproduction process.

A group of ten Rosella breeders will offer ten different methods for successful results. In truth, there is not one secret to success. Meetings are necessary for us all to provide the opportunity to exchange ideas. We cannot read enough. However, I suggest that we spend as much time as possible quietly observing our birds. By doing this, we will often detect things that make them feel uncomfortable. We have all heard of the many instances involving an unproductive pair that upon being moved into a different flight or even sold, went straight to nest and successfully reared young.

Perhaps a bit more time spent watching them would have informed the owner of the source of the problem.

Rosellas are not complicated in their habits, so neither should their management be complicated.

FEEDING SOFTBILLS – AN HISTORICAL SYNOPSIS

By Arthur Douglas

IFCB 1978

By “softbill,” bird keepers mean any small bird that feeds primarily upon insects rather than seed. Many softbills also live largely on berries or other fruit. Some eat a certain amount of seed. Of course, many kinds of “hardbills” will live wholly or mainly on insects and fruit when they have the opportunity.

Seed-eating birds are relatively easy to feed. Insect-eaters have always been considered more difficult and troublesome. In temperate climates their natural food is scarce and hard to come by in wintertime and, therefore, a substitute for it must be found. Such a food is often called a successful “softfood” and to be successful needs to be readily acceptable to the birds, nutritionally adequate, free from injurious elements, accessible and reasonably convenient for the bird keeper.

A survey of the literature dealing with bird keeping at once reveals great diversity of opinion on what is the best way of feeding birds that are predominantly insectivorous in nature. At the same time it reveals the conservatism of some bird keepers in holding fast to archaic methods and recipes.

Bird keeping, including the maintenance of softbilled birds,
dates back at least to the time of the Roman Empire. The Roman writer, Varro, claimed that bird keeping in Rome was initiated by Marcus Laenius Strabo of Brindisi (1). There is reason to believe the art is much more ancient (2).

Varro describes large aviaries in which fieldfares and European Blackbirds were held in numbers and fattened on a paste of figs and meal (ficis et farre) until they were in demand for a banquet and could be sold at a good profit. Varro’s own splendid aviary was kept strictly for pleasure, and in it were nightingales and blackbirds, among others. We are not told how they were fed.

Pliny writes of a white nightingale bought at a fabulous price and given to Agrippina, the emperor’s consort, and also of nightingales, magpies, a starling and a thrush that had been taught to imitate human speech (3). He says nothing about their food.

In the sixteenth century, early ornithologists, including Gesner (4) and Aldrovandi (5) give some indication of how certain birds were fed, but at least one writer dedicated a whole book to the art of bird keeping, including the keeping of softbills. He is Cesare Mancini Romano (6). A rather bad piracy of his work in French is better known (7). Mancini tells us how to keep and feed nightingales, blackcap warblers, rock-thrushes, songthrushes, blackbirds, calandras and skylarks as well as something he calls a “birdling”– perhaps a pipit.

Mancini advocates two alternative methods of feeding.

The first method of feeding is of raw sheep’s heart, cleared of fat and tough strings and finely chopped and ground. Yolk of hard-boiled egg can be substituted occasionally when fresh heart is not available. By way of a purgative or medicine, softbills should be given “those worms found in pigeon-lofts and also in flour”–presumably mealworms.

The second method of feeding is on a dry “paste”, which Mancini says is “marvelous” for softbills. He gives rather confusing instructions for making it.

If Mancini’s instructions in the sixteenth century are obscure, Oliva, in the succeeding century makes up for it. He gives a whole page of instructions and provides a splendid whole-page engraving showing the work in progress (8). This is what he advocates:

Peel half a pound of almonds. Break and grind them up fine. Stir into them four ounces of butter and the yolks of four hard-boiled eggs. Work this mixture into two or three pounds of chickpea flour. Put all this into a pan over a charcoal fire and keep stirring it until it appears to be fairly cooked. In a separate small pan melt and boil together a pound of honey and three ounces of butter. Remove any scum. Have one person to dribble the hot honey gradually through a perforated ladle onto the other ingredients while another person stirs constantly until the honey is well incorporated, when the paste will become granular. This is for summer use.

For winter use, add a half-pennyworth of saffron.

When the paste has become granular and is yellow in color, work it through a colander with round holes the size of vetch-seeds. Spread the grains on a white cloth to dry. Store them in a tin can or box.

The dry paste will keep for up to six months. If it becomes too dry, add a little honey to soften it.

R. L. Wallace was still recommending a paste almost identical to this at the end of the last century (9). A cheap and degenerate descendant of the original was available commercially in Wallace’s day under the name of “German paste”. It sold at three pennies a pound or less and usually consisted of pea-flour cooked with a little fat, sweetened with treacle or molasses and sprinkled with maw (poppy-seed) and hemp-seed. Superior samples had a few dried ants’ eggs and were then sold as “insectile food” (10).

German paste came under severe criticism among the more progressive fanciers of the 1900s. The Rev, C. D. Farrar had German paste in mind when he wrote “If you intend to feed only on peameal, I have done with you.” (11)

Despite its bad reputation, German paste seems to have been a fairly adequate staple diet for skylarks (12).

An Anonymous writer of 1697 continues to recommend the raw heart and the paste for feeding nightingales, but says he himself gives to his nightingales ground almonds, cake (massepain), chopped cabbage, lettuce and chickweed, cooked mussels, beets, peas and beans, sugar and other items (13).

By 1738, Eleazar Albin is recommending a mixture of the heart and hard-boiled egg as food for small softbills (14). The anonymous author of The Bird-Fancier’s Necessary Companion and Sure Guide does likewise (15).

Raw meat and egg remained popular through much of the last century. In 1826, the Rev. William Floyer Cornish of Totness in England claimed that he had been successful in keeping whitethroats and other small softbills on “beef, veal, mutton or lamb, not over-dressed, cut very small and mixed with hard eggs, yellow as well as white, and a little chopped hempseed, on which they have thriven very well” (16).

Similarly, in 1832, a Mr. Cox exhibited a nightingale in full song to members of the Zoological Society in London. He had kept it for four years. In the Arcana of Science, Mr. Cox expressed the opinion that failure with nightingales “and other Silviadae (sic)” was often due to over-elaborate feeding. He used finely ground meat and hard-boiled egg and considered insects by no means necessary (17).

Hemp-seed was recommended as an item of softbill diet by some bird keepers at an early date. Our nightingale author of 1697 condemns it, blaming it for causing fits in nightingales and starlings. On the other hand, Robert Sweet recommends for British warblers a staple diet consisting of scalded and finely “bruised” hemp-seed mixed to a moist paste with soft bread. He considers insects, fresh or dry, to be essential extras and accepts meat and egg as a useful occasional change. He observes that limy grit is essential and that the birds will not long stay healthy without it (18).

A contributor to the English journal Cage Birds was still advocating the use of crushed hemp-seed and bread in 1960. He added grated cheese. He mentions wrens, thrushes, blackbirds, larks and meadow pipits, and seems to consider it a recommendation for his food that birds fed on it develop white feathers in their wings and tails (19).

Dr. Bechstein, like Robert Sweet, recommends the use of fresh and dried insects, and in his Chamber-Birds gives instructions for laying in a stock for winter feed. However, his birds were largely fed on a “universal food” consisting of milksop mixed with either coarse wheatmeal or barleymeal. Alternatively, grated carrot could be mixed with the bread in place of the milk. Nightingales were given ants’ eggs, elderberries and cooked beefheart (20).

Dr. Bechstein set the style for softbill feeding in nineteenth century Europe. His universal foods may appear to us to be excessively austere. It would seem that some bird keepers of the last century saw things in a different light. In 1873 Swaysland considered oatmeal or barleymeal damped with milk to be the ideal diet for starlings, thrushes and blackbirds. For “warblers” (which in his day included chats and other small thrushes) he was still with Mancini in the sixteenth century and with Sweet in the early nineteenth (21).

Sumner Birchley was still using and promoting the use of dampened barleymeal for thrushes and starlings in 1909 (22). The birds were occasionally given a small earthworm or a snail “as a tit-bit”.

Apparently, some softbills were kept on what must surely be the ultimate in prison diets — wheat bran dampened with water. Field-fares seem to have survived on it. Bechstein says blackbirds will not. In our own century, Konrad Lorenz was still recommending it for starlings (23).

Since egg yolk has been a favorite bird food from time immemorial, it must have required moral courage on the part of Dr. George W. Creswel’ to condemn it roundly in the years 1903 and soon after (24).

Dr. Arthur Butler, a Founder-Member of the Avicultural Society, vigorously took up the defense of egg, boiled and as dried egg yolk (25).

A bitter correspondence between the two doctors soon divided the members of the Avicultural Society and the Foreign Bird Club into hostile camps — “eggists” and “non-eggists”. The non-eggists had considerable influence. It was soon obvious that Dr. Creswell was far better informed than his rival.

Even non-eggists of this period often fed their softbills largely on “stale sponge cake” — stale because they bought it at a reduced price as a bakery shop leftover.

While the no-eggist contention was at its peak, protagonists of both sides sought support for their opinions from distant cultures, especially from the Orient where bird keeping was known to be an accomplished art.

They found that in India softbilled birds were fed on satoo — a paste made from gram (chickpea flour) and ghee (clarified butter). This was supplemented with grasshoppers, maggots and other insects (26).

In China the traditional softbill food was made from finely chopped mutton or other meat soaked in a little water for a few hours and then rolled in egg powder and soya flour. (To this day, soya flour mixed with dried insects is still the staple diet for Zosterops spp. Eds.)
Insects were only offered during the moulting season (27). Frank Finn claimed that, in his day, softbills were sent from China to India on a diet of small shelled millet and dried insects and that they arrived in very good condition (26).

In Japan softbills were fed on fishmeal made from a smoke-dried freshwater fish. This was mixed with varying amounts of equal parts of rice flour and rice bran. Chopped greenfood was provided (28).

Towards the end of the nineteenth century, expanding international trade made available to European bird keepers dried insects that could be used in making acceptable mixtures called “insectile” foods. These soon became the accepted norm as soft food for insectivorous birds.

They were expensive compared with the older-fashioned diets, but every. one agreed that they were more natural and, therefore, better (29). The Rev. C. D. Farrar wrote of his own rather typical insectile mixture, “On the mixture and a few mealworms daily any sort of bird will do well.” (30)

A typical insectile food would be made up of equal parts by weight of dried ants’ eggs, dried flies and meatmeal, with a quantity of biscuit meal equal in weight to the rest of the ingredients combined. Some bird keepers used dried and ground silkworm pupae. Others added dried egg yolk. Still others put in some powdered cuttlebone. The mixture was made crumbly rather than powdery by the addition of fat, honey, or both, and was usually moistened before

use with hot or cold water, milk, grated carrot, grated apple, chopped lettuce or mashed potato (31). Cheese, though an old-fashioned and unnatural item, retained its prestige as an additive to insectile mixtures. Its chief protagonist in England was P. F. W. Galloway (32).

Mealworms have been considered a suitable food for captive soft-bills at least since Mancini’s time in the sixteenth century. Like Mancini, many bird keepers have believed that they could easily be overdone and that an excess would lead to gout, fits, obesity and other disorders. At the same time, the benefit to the birds of adding a generous ration of live food to some of the traditional staple diets has led to the belief, which I have often heard expressed, that live food must contain some mysterious vital element necessary to the birds and lacking in dead soft food. It is only in recent times that the “richness” of mealworms has been seriously questioned and their nutritional limitations indicated (33).

In the last hundred years, mealworms, maggots and, to a lesser extent, cockroaches, have been the live foods that have been most relief upon by keepers of softbilled birds. Live ants’ eggs and wasp-grubs have been important in their season, as have miscellaneous insects taken in the field with a sweeping-net or by beating. In recent years, considerable progress has been made in culturing a variety of different insects in quantity (34) and in developing methods of harvesting honeybee larvae for bird food (35). House crickets and locusts are being raised commercially. Since many kinds of softbilled birds will refuse to feed anything but fresh live food to their nestlings, these developments are of great practical importance to the aviculturists.

The development of moth-traps having an ultraviolet light and an electric fan that draws in night-flying insects attracted to it has opened up unprecedented opportunities for gathering wild insects in quantity. I, myself, have sometimes taken a pint of Noctuid moths in one night and have sometimes had two or three gallons of moths on hand, frozen, ready for use as needed. I am indebted to Mr. Harry Halff of San Antonio for pointing out the value of this device.

Insectile mixtures are probably still the main standby for bird keepers who only maintain small collections. Those who have to cater for considerable numbers of softbills have been able to benefit from a development that has gone counter to the natural food tradition. This counter-movement has been made possible by progress in our understanding of the elements of avian nutrition. A name particularly associated with the application of these principles to the feeding of captive birds is that of Dr. H. L. Ratcliffe (36).

Dr. Ratcliffe and others have set out to devise a diet that will be readily acceptable to the birds, more than adequate nutritionally, free from objectionable elements and reasonably economical, available and convenient for the bird keeper. Instead of insects, these foods are compounded from ingredients known to have superior food value. considering the proportion, availability and quality of their proteins, carbohydrates, fats, vitamins and minerals, etc. They are rather readily modified according to local conditions. Local variations have given good results in Switzerland (37), Canada (38), Texas (39) and no doubt elsewhere.

The poultry industry has been revolutionized in recent decades and mass-produced standard foods having controlled nutrient values have been devised to suit precisely the needs of poultry at different ages and stages.

No comparable detailed investigation of the precise nutritional needs of softbills has been feasible. Nevertheless, various mass-produced “Mynah Foods” and “Mockingbird Foods” are on the market. It would seem to be rather risky to assume that every brand put up for sale at a commercially competitive price is suitable in itself for any except the least demanding of omnivorous softbills. However, the potential of these convenient and economical foods should not be underestimated. I have seen Red-billed Leiothrix, Blue-winged Sivas and an American Blue Jay kept in immaculate feather and apparently in perfect health on an exclusive diet of dry “gamebird-starter”. They compared very favorably with lots of specimens of the same species I have seen maintained on more conventional diets.

Perhaps some mass-produced bird foods are already ideal for any except the most specialized feeders among softbills. These birds will always be a separate problem. It is certain that practical and satisfactory diets for the majority of softbills are now easily compounded by any bird keeper, diets that are far superior to those on which birds have been kept with at least some degree of success in the past. The day of insectile mixtures is probably past, and live food nowadays may be regarded as an expensive item reserved for birds that are newly acquired and are unused to any other food, or for breeding birds that are feeding young.

REFERENCES
III 1, Varro, Marcus Terentius 116-27 BC. Rerum Rusticarum, Book
2. Roberts, Sonia 1972. Bird-Keeping and Birdcages- A history, David + Charles, Newton Abbot 14.
3. Pliny the Elder. 23 or 24-79 AD. Historiae Naturalis, Book X.
4. Gesner, Conrad 1555. De Avibus.
5, Aldrovandi, Ulisse 1522-1605. Ornithologiae,
6, Mancini Romano, Cesare. Milan 1575 and 1645. Venice 1669. Ammaestramenti per allevare, pascere + curare gli uccelli.
7. Instructions pour elever, nourrir, instruir et penser toutes sortes de petits oyseaux de voliere. Paris 1647 and 1697.
8. Olina, Gio, Pietro 1684. Uccelliera, Rome,
9, Wallace, R.L. 1889. British Cage Birds, 189, London.
10. Frostick, John 1903. Bird Notes, Vol. 1, 124, Brighton.
11. Farrar, C.D. about 1920. Through a Birdroom Window, 33. London.
12. The Skylark, Nutshell Series No. 3. Cagebirds London 1909.
13. Traite du Rossignol 1697. 59. Paris.
14. Albin, Eleazar 1738. A Natural History of English Song Birds, 71. London.
15. The Bird Fancier’s Necessary Companion and Sure Guide. 1762. 14, London.
16. Bewick, Thomas. 1832. A History of British Birds, I, footnote p. XVI. Newcastle.
y I
17. Goldsmith’s Natural History. Edited by Henry Innes, footnote, 139.
18. Sweet, Robert. 1823. The British Warblers, 2, London.
19. Wilkins, A.J. 1960. Cage Birds, 22. London,
20. Bechstein, Dr. J.M. 1890. The Natural History of Cage Birds, 10. London.
21. Swaysland, W. 1873. The Illustrated Book of Cage Birds, 299, London.
22. Birchley, Summer W. 1909. British Birds for Cages, Aviaries and Exhibition, II, 71, London.
23. Lorenz, Konrad Z. 1955. King Solomon’s Ring, 61, London.
I I
24. Creswell, W. George 1904. Bird Notes, Brighton.
25, Aviculture Magazine I, New Series, 1903,
Butler, Dr. A.G. 1904, Aviculture Magazine II, New Series,
26. Finn, Frank 1901. The Cage Birds of Calcutta, The Ibis
27. Delacour, Jean 1928. Avicultural Magazine, Fourth Series VI, 29.
28. Taka-Tsurkasa, N. 1921. Avicultural Magazine, Third Series, XIII, 49
29. Frostick, John 1903. Bird-Notes I, 121.
30. Farrar, C.D. 1920. Through a Bird-Room Window, 35.
31. Fulljames, H.J. 1910. Insectivorour British Birds, Nutshell Series, nos 25 and 26, London.
32. Galloway, P.F.M. 1925. How to keep insectivorous Birds in Perfect Condition. Avicultural Magazine, Fourth Series, III, 46.
33. Roots, Clive 1966. Maggots and Mealworms, Avicultural Magazine, LXXIV, 186
34. Meadon, F. 1961. Live Food for Birds, Cage & Aviary Birds, London.
35. Ficken, R.W. and W.C. Dilger 1961. Insects and Food Mixtures for Insectivorous Birds. Avicultural Magazine, LXVII, 27.
36. Ratcliffe, Herbert L. 1966. Diets for Zoological Gardens, International Zoo Yearbook, VI, 4, London.
37. Wackernagel, Hans 1966. Feeding Wild Animals in Zoological Gardens, International Zoo Yearbook, VI, 23, London
38. Ivor, Hance Roy 1964. Personal Communication, Ontario.
39. Douglas, A. 1971. Soft-food of Outstanding Merit, Cage and Aviary Birds, 7.

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