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Footedness in parrots: three centuries of research, theory, and mere surmise.

In contemporary debates about laterality in animals, the parrot (Psittaciformes) is often cited as an exemplar--possibly unique--of laterality in limb function at the population level comparable in kind and strength to handedness in man. This conclusion rests on just two reports (Friedman & Davis, 1938; Rogers, 1980) that most species of parrots are left-footed, that is, that they preferentially perch on the right foot and hold food with the left. In fact, speculation about and scientific study of laterality in parrots go well beyond these two investigations. The question itself dates from at least the 17th century, after which, beginning in the 1860s, it became the subject of broad interest and debate. In our own time, it also has continued to occupy the attention of at least a small number of ornithologists and field biologists whose work, like that of their predecessors, is not cited in the current neuropsychological literature on this topic. To fill out the historical as well as contemporary record for consideration by neuropsychologists today, these other observations and theoretical analyses are reviewed, and new questions about laterality in parrots raised by this work are presented.

Animals↗

Redescription of Alcedinectes alcyon (Acari:Hypoderatidae) from the belted kingfisher (Aves:Coraciiformes; Alcedinidae).

Alcedinectes alcyon (Boyd) is redescribed based on specimens collected from its belted kingfisher, Megaceryle alcyon (L.), type host in Florida. This species differs from A. cerylei Fain by the chaetotaxy of tarsus III. A. alcyon has a long stout apical seta that tapers to a curved tip resembling that seen on tarsus III of the genus Tytodectes (Ispidectes) Fain. The equivalent seta in A. cerylei is a stout spine. There are other minor differences between these species in the pattern of idiosomal sclerotization and leg chaetotaxy. The chaetotaxy and solenidiotaxy of legs I and II in Alcedinectes closely resemble that of Amazonectes psittaci Fain & Vercammen-Grandjean from parrots (Psittaciformes), whereas the morphology and chaetotaxy of the idiosoma resemble that of Tytonectes (Ispidectes) spp. which also occurs in kingfishers (Coraciiformes). The long stout seta that tapers to a curved tip on tarsus III in A. alcyon also links the genus Alcedinectes with Tytodectes (Ispidectes).

Animals↗

Complex distribution of avian color vision systems revealed by sequencing the SWS1 opsin from total DNA.

To gain insights into the evolution and ecology of visually acute animals such as birds, biologists often need to understand how these animals perceive colors. This poses a problem, since the human eye is of a different design than that of most other animals. The standard solution is to examine the spectral sensitivity properties of animal retinas through microspectophotometry-a procedure that is rather complicated and therefore only has allowed examinations of a limited number of species to date. We have developed a faster and simpler molecular method, which can be used to estimate the color sensitivities of a bird by sequencing a part of the gene coding for the ultraviolet or violet absorbing opsin in the avian retina. With our method, there is no need to sacrifice the animal, and it thereby facilitates large screenings, including rare and endangered species beyond the reach of microspectrophotometry. Color vision in birds may be categorized into two classes: one with a short-wavelength sensitivity biased toward violet (VS) and the other biased toward ultraviolet (UVS). Using our method on 45 species from 35 families, we demonstrate that the distribution of avian color vision is more complex than has previously been shown. Our data support VS as the ancestral state in birds and show that UVS has evolved independently at least four times. We found species with the UVS type of color vision in the orders Psittaciformes and Passeriformes, in agreement with previous findings. However, species within the families Corvidae and Tyrannidae did not share this character with other passeriforms. We also found UVS type species within the Laridae and Struthionidae families. Raptors (Accipitridae and Falconidae) are of the violet type, giving them a vision system different from their passeriform prey. Intriguing effects on the evolution of color signals can be expected from interactions between predators and prey. Such interactions may explain the presence of UVS in Laridae and Passeriformes.

Amino Acid Sequence↗

Ultraviolet signals in birds are special.

Recent behavioural experiments have shown that birds use ultraviolet (UV)-reflective and fluorescent plumage as cues in mate choice. It remains controversial, however, whether such UV signals play a special role in sexual communication, or whether they are part of general plumage coloration. We use a comparative approach to test for a general association between sexual signalling and either UV-reflective or fluorescent plumage. Among the species surveyed, 72% have UV colours and there is a significant positive association between UV reflectance and courtship displays. Among parrots (Psittaciformes), 68% of surveyed species have fluorescent plumage, and again there is a strong positive association between courtship displays and fluorescence. These associations are not artefacts of the plumage used in courtship displays, being generally more 'colourful' because there is no association between display and colours lacking UV reflectance or fluorescence. Equally, these associations are not phylogenetic artefacts because all results remain unchanged when families or genera, rather than species, are used as independent data points. We also find that, in parrots, fluorescent plumage is usually found adjacent to UV-reflective plumage. Using a simple visual model to examine one parrot, the budgerigar Melopsittacus undulatus, we show that the juxtaposition of UV-reflective and fluorescent plumage leads to a 25-fold increase in chromatic contrast to the budgerigar's visual system. Taken together, these results suggest that signals based on UV contrast are of special importance in the context of active sexual displays. We review briefly six hypotheses on why this may be the case: suitability for short-range signalling; high contrast with backgrounds; invisibility to predators; exploitation of pre-existing sensory biases; advertisement of feather structure; and amplification of behavioural signals.

Animal Communication↗

Reclassification of Bisgaard taxon 33, with proposal of Volucribacter psittacicida gen. nov., sp. nov. and Volucribacter amazonae sp. nov. as new members of the Pasteurellaceae.

A total of 25 strains isolated from parrots, budgerigars, parakeets (Psittaciformes) and a chicken, mostly associated with respiratory disease or septicaemia, were classified as a new genus, Volucribacter gen. nov., within the family Pasteurellaceae, on the basis on unique phenotypic characteristics and clear monophyly as determined by 16S rRNA gene sequence comparison. Comparison of 16S rRNA gene sequences from six strains showed at least 98.8 % similarity and the closest similarity outside the genus was found to Bisgaard taxon 34 and to Pasteurella avium, at 94.6 and 94.5 %, respectively. Phenotypes that separate the new genus from other genera of the Pasteurellaceae included at least two characters. The genus includes two species, Volucribacter psittacicida sp. nov. and Volucribacter amazonae sp. nov., corresponding to the two biovars previously outlined, underlining that most isolates have been obtained from psittacine birds. The two species can be separated by fermentation of meso-inositol, (-)-L-fucose, maltose and dextrin and a positive ONPG test. The type strains for Volucribacter psittacicida and Volucribacter amazonae are respectively Gerl. 236/81(T) (=CCUG 47536(T)=DSM 15534(T)) and 146/S8/89(T) (=CCUG 47537(T)=DSM 15535(T)).

Animals↗

Description of Sarcocystis falcatula stiles, 1893, a parasite of birds and opossums.

Sarcocystis falcatula Stiles, 1893 is re-described. Intermediate hosts of the parasite which was earlier described as Sarcocystis debonei Vogelsang, 1929 are species of passeriform, psittaciform, and columbiform birds. In these birds, muscle zoites are 6.88 X 2.19 (4.8-8.4 X 1.2-3.6) micron and are enclosed in a cyst wall with regular protrusions, 1-5 micron long. The convoluted primary wall has multiple thin areas in the osmiophilic layer. Microtubules originate in the ground substance and extend to the tips of the protrusions. The only known definitive host is the opossum, Didelphis virginiana; rats, cats, a dog, and a ferret could not be infected from muscle cysts. Sporocysts from opossums infected from five different infected avian sources measure 11.2 X 7.4 (9.6-12.0 X 6.0-8.4) micron.

Animals↗

Guidelines for the interpretation of laboratory findings in birds and mammals with unknown reference ranges: plasma biochemistry.

The results of analyses of plasma samples from healthy and sick adult birds and mammals of a wide range of species were used to calculate reference ranges and clinical ranges for 11 biochemical variables (calcium, sodium, chloride, total protein, albumin, globulin, inorganic phosphate, potassium, bicarbonate, urea and creatinine concentrations) in six mammalian Orders (Primata, Carnivora, Pinnipedia, Proboscidea, Perissodactyla and Artiodactyla) and six avian Orders (Ciconiiformes, Gruiformes, Pelecaniformes, Psittaciformes, Falconiformes and Strigiformes). The patterns of variation between Orders for the different biochemical variables were then used to provide guidelines for the interpretation of plasma biochemistry results in the species in these Orders for which there are no reference data. The observed clinical ranges provide an indication of the potential diagnostic value of each variable.

Animal Diseases↗

Comparative distribution of 2[125I]iodomelatonin binding in the brains of diurnal birds: outgroup analysis with turtles.

The roles that the pineal gland and its hormone melatonin play in the regulation of circadian rhythmicity and photoperiodism vary among vertebrate species. Recently, putative sites of melatonin action have been elucidated in several avian and mammalian species by application of in vitro binding of a radioiodinated melatonin agonist, 2[125I]iodomelatonin (IMEL) and autoradioradiography. These studies in mammals, birds and reptiles have indicated profound differences in the distribution of IMEL binding between these diverse groups, suggesting that these large differences in binding may reflect differences in melatonin function. The present study was performed to determine systematically whether the variance in IMEL binding among avian species corresponds to changes in circadian organization and/or phylogenetic relationships. The distribution of specific IMEL binding was determined in the brains from birds belonging to 14 different species in 5 Orders (Psittaciformes, Passeriformes, Columbiformes, Galliformes and Anseriformes) using in vitro binding, autoradiography and computer-assisted image analysis. The distribution was compared to a similar study in 3 species of turtles as an outgroup. The data indicated IMEL binding in retinorecipient structures of the circadian, tectofugal, thalamofugal and accessory optic visual pathways in all avian species. Relay nuclei and integrative structures of the tectofugal, thalamofugal, accessory optic, and limbic systems, however, bound the hormone to varying degrees. In turtles, binding was observed in retinorecipient structures of the thalamofugal visual pathway and in retinorecipient and integrative areas of the tectofugal visual pathway. No binding was observed in the pineal gland, tuberal hypothalamus or adenohypophysis in any avian or testudine species. This distribution is drastically different from that observed in mammals, where binding predominates in the pars tuberalis of the adenohypophysis and in the suprachiasmatic nucleus, suggesting that the circadian system may influence a wide array of sensory and integrative functions in birds and reptiles through the circadian secretion of melatonin, but that this capacity has been lost in mammals.

Animals↗

Avian papillomaviruses: the parrot Psittacus erithacus papillomavirus (PePV) genome has a unique organization of the early protein region and is phylogenetically related to the chaffinch papillomavirus.

BACKGROUND: An avian papillomavirus genome has been cloned from a cutaneous exophytic papilloma from an African grey parrot (Psittacus erithacus). The nucleotide sequence, genome organization, and phylogenetic position of the Psittacus erithacus papillomavirus (PePV) were determined. This PePV sequence represents the first complete avian papillomavirus genome defined. RESULTS: The PePV genome (7304 basepairs) differs from other papillomaviruses, in that it has a unique organization of the early protein region lacking classical E6 and E7 open reading frames. Phylogenetic comparison of the PePV sequence with partial E1 and L1 sequences of the chaffinch (Fringilla coelebs) papillomavirus (FPV) reveals that these two avian papillomaviruses form a monophyletic cluster with a common branch that originates near the unresolved center of the papillomavirus evolutionary tree. CONCLUSIONS: The PePV genome has a unique layout of the early protein region which represents a novel prototypic genomic organization for avian papillomaviruses. The close relationship between PePV and FPV, and between their Psittaciformes and Passeriformes hosts, supports the hypothesis that papillomaviruses have co-evolved and speciated together with their host species throughout evolution.

5' Untranslated Regions↗

Developmental patterns of heart rate in altricial avian embryos and hatchlings

The aims of this study were to determine the patterns of development of heart rate (fH) in altricial avian embryos and hatchlings, and then to examine how fH is regulated to meet metabolic requirements in altricial embryos. Embryonic mean heart rate (fH-) in 12 altricial species (Passeriformes and Psittaciformes) increased during pre-pipping incubation in all species except the cockatiel (Nymphicus hollandicus), in which fH- tended to decrease prior to pipping. The rate of increase in fH- tripled during the pipping phase in all species, and fH- was significantly higher during the pipping period and in hatchlings than during pre-pipping development. The O2 pulse (O2 consumed per cardiac beat) of altricial embryos increased in direct proportion to embryo mass (loge/loge base), although fH- was often low prior to pipping, implying that stroke volume increases in the second half of incubation. We conclude that fH contributes more than other factors towards supplying the metabolic demands of the embryo during the middle of incubation and the final pipping phase, but less during the intervening period of late incubation.

Journal Article↗

The skull development of parrots with special reference to the emergence of a morphologically unique cranio-facial hinge.

The order Psittaciformes (parrots) has unique morphological features in the head that are evolutionarily novel. To better understand the unique evolution of the head in parrots, the developmental pattern of the skull of the budgerigar (Melopsittacus undulatus) was initially described on the basis of transparent skeletal specimens. Although the fundamental pattern of the skull development of birds is conserved in parrots, some differences were observed between parrots and other groups of birds. In parrots, the vacuity in the interorbital septum did not emerge throughout ontogeny, in contrast to other lineages of birds, for example Galliformes and Coliiformes. This feature seems to be concerned with the attachment of the unique jaw muscle of parrots, M. ethmomandibularis, to the interorbital septum. In spite of a prokinetic skull, the cranio-facial hinge of parrots was brought about by secondary transformation of dermal bones unlike that of birds with a standard prokinetic skull (e.g. Corvus) in which the nasal-frontal suture directly becomes a hinge of bending. To further understand the evolution of "pseudoprokinesis" in parrots, the construction of a robust avian phylogeny is desired. The parrot-specific suborbital arch and cranio-facial hinge are not seen until birds leave the nest and can feed themselves. In conclusion, these structures are considered to be essential for eating hard and/or large meals.

Adaptation, Physiological↗

Dispharynxiasis in a captive princess parrot.

The acuariid nematode Dispharynx nasuta was found in a princess parrot, Polytelis alexandrae, at the National Aviary in Pittsburgh (Pennsylvania, USA). This is the first report of D. nasuta from the host order Psittaciformes, and was the presumed cause of death in this parrot.

Animals↗

[Polyomavirus infections in exotic birds in Switzerland].

Polyomavirus infections in budgerigars, in other parrots as well as in passeriformes have been described in many countries, but not in Switzerland so far. This paper reports on cases of polyomavirus infections in three different bird collections in Switzerland. The first outbreak occurred in a mixed collection of passeriformes, the second in a collection of parrotlets (Genus Forpus), and the third in a large groups of parrots (Psittaciformes), of which only one recently acquired Kea (Nestor notabilis) was affected. The clinical symptoms, gross necropsy changes and histopathologic findings are described. The presence of avian polyomavirus was confirmed by the polymerase chain reaction (PCR).

Animals↗

Avian influenza A viruses in birds --an ecological, ornithological and virological view.

Avian influenza A viruses (AIV) are the causative agents of the presently most important poultry disease. Ten countries in Asia and several other countries in Eastern Europe suffer high losses from the lethal effects of these viruses of the H5N1 subtype. AIV of other subtypes cause in additional countries severe losses. The threat to health and well-being of the avifauna, domestic poultry and possibly mammals including humans are worldwide of major concern. The European Union reacted with a complete import ban on untreated meat, eggs, poultry products as well as free-living and pet birds. Extensive surveillance of free-living birds and domestic poultry that is maintained in free-range and close to open waters were initiated in an attempt to gather information on the current status of infection with these viruses and to target appropriate countermeasures for the protection of domestic poultry (in-house keeping) and to safeguard food production for humans. Since the monitoring of free-living birds is labour-intensive, costly, and time-consuming, only birds should be included in the monitoring programme that harboured in the past most if not all influenza A viruses. The birds of the order Anatiformes, family Anatidae, subfamilies Anserinae and Anatinae, provided 65.9 % of all avian AIV isolates. The cosmopolitan Common Mallard (Anas platyrhynchos) is the dominant species with the highest rate of isolations among all bird species. Second in frequency is the North-American Blue-winged Teal (Spatula discors). Consequently, free-living anatiform birds of the genera Anas and Spatula should comprise the main focus for the collection of cloacal and pharyngeal swabs. With the likely exception of the most recent H5N1 viruses, signs of disease were not recorded in AIV infected anatiform birds. AIV isolations were definitely less frequently obtained from birds of the orders Phasianiformes (including domestic chickens and turkeys), Charadriiformes (plovers and lapwings), Lariformes (gulls), Columbiformes (pigeons) and Psittaciformes (psittacines) and need less attention in sampling efforts. This review presents also data on taxonomy and most suitable means for isolation and typing of haemagglutinating viruses. The different frequencies of the detection of 16 haemagglutinin (HA) subtypes and 9 subtypes of neuraminidase (NA) surface antigens are composed on the basis of extensive literature retrievals. Both antigens occure in isolates at different frequencies. Only 103 of all 144 possible HA x NA combinations were described so far. The AIV that contain the HA subtypes H3, H4, H6 are most frequently isolated whereas the AIV of the subtypes H5 and H7 were less frequently encountered. All other HAs are rather rare. AIV that possess the NA of the subtypes N2, N1, N8 and N3 are frequent and all other NAs are rarely detected.

Animals↗

Light microscopic morphometry of the kidneys of fourteen avian species.

Kidney volume and the volume proportions of the cortex, medulla, blood vessels larger than capillaries and ureter and ureteral ducts were investigated in 61 birds from 14 species representing passeriformes, psittaciformes, podicipediformes, anseriformes, galliformes, and columbiformes. The kidneys were fixed in situ by perfusion via their arterial supply. Kidney volume and body weight were strongly correlated (r = 0.991); the allometric function relating kidney volume (Vk) to body weight (W) was: Vk = 13.96W0.902. Histological sections of one kidney from each bird were analysed by point counting with a Zeiss integrating eyepiece to estimate the volume proportions and subsequently the absolute volumes of the main components of the kidney. The avian kidney was found to consist of a very large cortex (range 71-81%), a relatively small medulla (range 5-15%), blood vessels larger than capillaries (range 10-13%). For all the birds, the mean volume proportions of the kidney components were as follows: cortex, 77%; medulla, 10%; blood vessels larger than capillaries, 12%; ureter and ureteral ducts, 1%. The values for the absolute volume of each of the main components of the kidney were well correlated with body weight. The allometric equations and correlation coefficients (r) relating: (a) the volume of cortex (Vc) and the body weight (W) was Vc = 10.06W0.914, r = 0.991; (b) the volume of medulla (Vm) and body weight was Vm = 2.27W0.795, r = 0.974; (c) the volume of blood vessels larger than capillaries (Vb) and body weight was Vb = 1.76W0.894, r = 0.988. All the birds were divided into two groups in the following five sets: (a) passeriform and non-passeriform orders; (b) galliform and non-galliform orders; (c) small and large birds weighing less than and more than 500 g respectively; (d) aquatic and non-aquatic birds; (e) desert and non-desert birds. In the two groups of each set the differences between kidney volume per gram body weight and those between the respective percentage proportions of cortex, medulla and blood vessels larger than capillaries were assessed by Student's t-test. The differences between kidney volume per gram body weight were significant for all sets except desert and non-desert birds; those between the respective percentage proportions of cortex and medulla were significant for all sets; those between the percentage proportions of blood vessels larger than capillaries were significant only in desert and non-desert birds.(ABSTRACT TRUNCATED AT 400 WORDS)

Anatomy, Comparative↗

The intermediate host spectrum in a Sarcocystis species of birds.

The life cycle of an avian Sarcocystis has been completed in the laboratory, originating with naturally infected icterids and passing alternately between opossums (Didelphis virginiana) and experimentally infected birds. To determine the intermediate host range, six avian species, including canaries (Serinus canarius), zebra finches (Poephila guttata), budgerigars (Melopsittacus undulatus), pigeons (Columba livia), chickens (Gallus gallus), and guinea fowl (Numida meleagris), were inoculated orally with Sarcocystis sporocysts derived from experimentally infected opossums. All birds but the Galliformes were susceptible to merogony. Pigeons (Columbiformes) were susceptible to early merogony but apparently not to muscle stages. Passeriformes and Psittaciformes were completely susceptible and the parasite developed into muscle cysts in them.

Animals↗

Allometric comparison of brain and main brain subdivisions in birds.

The brain weights and volumes of the main brain subdivisions: telencephalon, diencephalon, mesencephalic tegmentum, optic tectum, cerebellum and myelencephalon were measured in 8 species of Phasianoidea (Galliformes) and in 20 species of birds belonging to 13 orders. The sizes of these structures were compared using allometry formulae calculated for each structure from the 8 Phasianoidea. Encephalization and the relative size of the main brain subdivisions are expressed by size indices. Higher encephalization indices are obtained in species which are either highly telencephalized as in Psittaciformes or with a large cerebellum as in Sphenisciformes. The brainstem components: diencephalon, mesencephalic tegmentum and myelencephalon appear to be more conservative in their size variation than the telencephalon, cerebellum and optic tectum. The size of the cerebellum appears to be related to flight abilities. The size of the optic tectum is related to the importance of vision in foraging within the Charadriiformes. In tactile-feeding Charadriiformes and filter-feeding Anseriformes, the optic tectum is relatively small.

Animals↗

[The modified Schirmer tear test in birds--a method for checking lacrimal gland function].

The Schirmer tear test (STT) is an important component of the complete ophthalmic examination in mammals for evaluating lacrimal function. However, no STT reference values for birds have been published to date. 255 ophthalmologically healthy birds (42 different species form seven orders) were tested using a 1-minute STT I (without topical anaesthesia, reflex secretion), STT II (with topical anaesthesia, basal secretion) and as a modification STT III (measurement under isofluran/oxygen, inhalation anaesthesia, a precaution to minimize examination induced stress) using standardized 5-mm-STT strips as well as--under consideration of the size of the palpebral fissure--strips trimmed down to 2 and 4 mm. Mean values (+/- SD) for STT I and STT II were 3.2 +/- 2.7 to 7.5 +/- 2.6 mm/min and 1.7 +/- 1.4 to 4.5 +/- 2.8 mm/min respectively in Psittaciformes (using 4 mm strips), 4.1 +/- 2.7 to 14.4 +/- 7.2 mm/min and 2.0 +/- 1.7 to 4.2 +/- 3.1 mm/min in Falconiformes and 10.7 +/- 4.0 to 11.5 +/- 5.4 mm/min and 3.6 +/- 1.7 to 5.9 +/- 3.1 mm/min in Accipitriformes (using 5 mm strips). Due to the lack of a glandula lacrimalis Strigiformes showed STT levels below 3 mm/min. STT III values were between those of STT I and STT II in Falconiformes and Accipitriformes but not in Columbiformes. On the basis of these results the STT will become an important component of the bird specific ophthalmologic examination procedure allowing more sophisticated examination of lacrimal gland associated avian eye disorders.

Animals↗