Diet and Geophagy Across a Western Amazonian Parrot Assemblage

We identified species‐ and community‐level dietary characteristics for a species‐rich Amazonian parrot assemblage to determine relationships among dietary metrics and use of geophagy sites. Previous studies suggest that soil is consumed at geophagy sites in this region mainly to supplement dietary sodium. We accumulated 1400 feeding records for 16 parrot species over 2 yr and found that seeds, flowers, and fruit pulp featured prominently in diets, while bark, insects, and lichen were consumed in small quantities. Food availability across 1819 trees was measured, and we found that flower availability was highest in the dry season and fruit production peaked in the wet season, but that phenology patterns of the 20 most commonly foraged plant species suggest no serious food bottlenecks. Partitioning of available food resources among the 13 most commonly encountered parrots is suggested by an ordination analysis (DCA), which placed the large macaws (Ara) with the Amazona parrots at the ‘primary forest’ end of a dietary resource axis and four smaller species at the ‘successional forest’ end of the axis. Parrot species associated with successional forest also consumed less plant species overall. Furthermore, these parrot species consuming successional forest resources had higher claylick visitation rates than those consuming primary forest resources suggesting they derive the greatest benefits from soil consumption.     

Insights into the feeding ecology of the Seychelles Black Parrot Coracopsis barklyi using two monitoring approaches

Feeding ecology is an important factor for the survival of a species and knowledge of its parameters is a prerequisite for successful conservation work. In this study we describe the feeding ecology of the endemic Seychelles Black Parrot Coracopsis barklyi on Praslin, Seychelles, the only island on which this parrot is resident. We compared two methods to evaluate feeding choices: incidental observations and feeding walks on 25 transects in all habitat types. Black parrots fed on 46 different species, bringing the total number of known food plants to 53 species. They predominantly consumed endemic and native species (58% of observed feeding bouts), mainly their fruit pulp (in 68% of feeding bouts), followed by buds (15%) and seeds (37%) with occasional observations of leaves, bark and scale insects. The incidental method rendered many more observed bouts than the transect approach and the ratios of consumed species differed between methods but the transect results are regarded as more representative. The incidental method is not suitable for quantitative conclusions but complements the transect method, providing information about rarely occurring feeding events.     

Lust for Salt in the Western Amazon

Although the use of mineral licks by diverse Amazonian birds and mammals is well‐known, the ultimate motivation for such behavior remains unclear. As aerosol deposition of salts declines with distance from oceanic sources, lick visitation in the western Amazon can be explained by demand for sodium, given the low concentration of this micronutrient in the plant tissues consumed by these taxa. Sodium limitation also influences ant foraging behavior, and impinges on ecosystem rates of carbon cycling. The biogeographical context of sodium availability has been largely overlooked, but has substantial pantropical implications for herbivore and decomposer performance in inland rain forests. Abstract in Spanish is available in the online version of this article.     

A molecular phylogeny of Amazona: implications for Neotropical parrot biogeography, taxonomy, and conservation

Amazon parrots (Genus Amazona) are among the most recognizable and imperiled of all birds. Several hypotheses regarding the evolutionary history of Amazona are investigated using a combined phylogenetic analysis of DNA sequence data from six partitions including mitochondrial (COI, 12S, and 16S) and nuclear (beta-fibint7, RP40, and TROP) regions. The results demonstrate that Amazona is not monophyletic with respect to the placement of the Yellow-faced parrot (Amazona xanthops), as first implied by. In addition, the analysis corroborates previous studies suggesting a Neotropical short-tailed parrot genus as sister to Amazona. At a finer level, the phylogeny resolves the Greater Antillean endemic species as constituting a monophyletic group, including the Central American Amazona albifrons, while further revealing a paraphyletic history for the extant Amazon species of the Lesser Antilles. The reconstructed phylogeny provides further insights into the mainland sources of the Antillean Amazona, reveals areas of taxonomic uncertainty within the genus, and presents historical information that may be included in conservation priority-setting for Amazon parrots.     

Parrot behavior at a Rio Manu (Peru) clay lick: temporal patterns,associations, and antipredator responses

Although eating clay at licks (a form of geophagy) has been described, there are few behavioral data on temporal patterns, social interactions, species associations, or reactions to potential predators. We examined the behavior of nine species of macaws, parrots, and parakeets at the Machiguenga Ccolpa, a clay lick on the Rio Manu, Peru in the dry season. Three distinct mixed-species groups used the licks: in the early morning (parrots and small macaws), in mid-morning (large macaws), and in the early afternoon (parakeets), although the latter two groups used the licks at other times of day as well. The first parrots to begin eating at the lick in the early morning were yellow-crowned parrots (Amazona ochrocephala) and dusky-headed parakeets (Aratinga weddellii), followed by blue-headed parrots Pionus sordidus, and then by mealy (Amazona farinosa) and orange-cheeked (Pionopsitta barrabandi) parrots, and chestnut-fronted macaws (Ara severa). Although blue-headed parrots fed in dense groups of over 50, the others rarely exceeded 20 individuals. Scarlet macaws (A. macao) sometimes fed alone or joined the early morning groups, but most associated with a large group of red and green macaws (A. chloroptera) that arrived, often scaring off the smaller birds. On average, about 100 macaws and parrots fed in the early morning, macaw feeding groups averaging just over 40, and the parakeets averaged over 70. Average time at the lick ranged from 28 min for yellow-crowned parrots to 47 min for tui parakeets. Of the early morning group, blue-headed and mealy parrots were the most aggressive and orange-cheeked parrots were the least aggressive. Red and green macaws were more aggressive than scarlet macaws; the parakeets were equally aggressive. All species had more aggressive interactions with conspecifics than with other species. Responses to intruders and predators varied by species of parrot/macaw and type of intruder. In response to intruders or loud calls, responses could be partial (some individuals flew away, circled, and returned), temporary (all individuals flew away but returned within a few minutes), or total (all flew away and abandoned feeding for at least a half hour). The large macaws showed the lowest rate of total abandonment and the parakeets showed the highest. People passing up or down river in boats scared birds from the lick. The local residents (Machiguenga tribespeople in boats) elicited a much greater response than did the researchers. In the recent past, macaws and parrots were hunted for food, feathers, and the pet trade, and the birds response, as well as the presence of parrot and macaw feathers in local villages we visited, suggests some continued exploitation, or a long-term memory in the birds.Communicated by R.F. Oliveira     

A cytogenetic study of four species of cockatoos and Amazon parrots

The karyotypes of three species of cockatoos, Cacatua moluccensis, Cacatua goffini, and Cacatua sanguinea, and one Amazon parrot, Amazona aestiva, are described for the first time. These karyotypes are compared to the twenty-three species of psittacines published previously. Cacatua moluccensis appears to be a link in the evolutionary tree between Probosciger aterrimus and the other cockatoo species studied. This study reaffirms that the cockatoos are probably most closely related to the loris (Loriculus sp.) and Amazon parrots based on karyotype (M. W. M. Van Dongen & L. E. M. De Boer, (Genetica 65: 109–117) 1984).     

Parrot claylick distribution in South America: do patterns of “where” help answer the question “why”?

Geophagy is well known among some Neotropical parrots. The clay apparently adsorbs dietary toxins and/or provides supplemental nutrients. We used location data and 23 environmental layers to develop a predictive model of claylick distribution using Maxent software. We related species characteristics to claylick use and examined how parrot assemblages using claylicks changed with distance from the centre of claylick distribution. Fifty‐two parrot claylicks were reported from an area of ca 4 million km² but over 50% were restricted to a 35 000 km² region of southeast Peru and northern Bolivia. Claylicks were strongly associated with moist forest on younger (<65 millions of yr) geological formations and exposed river banks. The predictive model of claylick distribution matched our reported range well, with precipitation of warmest quarter, land cover, temperature seasonality, and distance from the ocean being most important predictors of claylick presence. Twenty‐six of the region's 46 parrot species visited claylicks. Species differed greatly in their lick use, but body size, dietary breadth, abundance and other traits were poor predictors of lick use. We are confident that our survey identified the distribution of major parrot claylicks in South America, although less conspicuous parrot geophagy may occur elsewhere. We suggest that claylick distribution reflects both underlying geology (allowing claylick formation in only some regions) and the physiological need for geophagy among parrots in different parts of the continent. Data on the latter are inconclusive, but we argue that parrot claylick distribution supports the contention that geophagy is related more to sodium deficiencies than to protection from dietary toxins.     

The sentineling—Foraging trade‐off in dominant and subordinate Arabian babblers

Many cooperative breeders forage under predation risks, sentineling is a central activity, and groupmates have to balance between sentineling and foraging. The optimal balance between sentinel activity and foraging may differ among dominant and subordinate individuals, as dominants are more efficient foragers. Two theoretical models pertain to this balance and predict when individuals with different foraging abilities should switch between the two activities on the basis of their energetic state. In one of these models, individuals must attain a critical energetic level by dusk to pass the night, and in the second model fitness is monotonically increasing with the energetic state. We tested these models in the cooperatively breeding Arabian babbler, Turdoides squamiceps. We measured the length of sentinel bouts and the gaps between them both in natural conditions and following experimental feeding. Following feeding ad libitum, subordinates expanded their sentinel bouts significantly more than dominants in comparison with natural conditions. These findings are consistent with the first model, but not with the second. In the experiment, we measured the mass of mealworms consumed by each individual following a sentinel bout relative to its body mass. This ratio was larger for subordinates, indicating that they ended their sentinel bouts at a lower energetic state than dominants. This finding is consistent with the second model, but not with the first. Immediately after eating ad libitum, in 62% of the cases the first behavior performed by the babblers was a new sentinel bout, but in 17% it was a mutual interaction with a groupmate, indicating that social interactions also play a role in the trade‐off vis‐à‐vis sentinel activity.     

Characterisation of 13 microsatellite loci for the Moluccan Cockatoo, Cacatua moluccensis, and Cuban Amazon, Amazona leucocephala, and their conservation and utility in other parrot species (Psittaciformes)

The parrot family consists of approximately 330 species, many of which are endangered due to habitat destruction and illegal trade. Microsatellite markers can provide important tools for examining both conservation and forensic issues in this family, and the availability of additional markers will prove useful for studies of other species in the parrot family. Parrots have proved a difficult family from which to develop microsatellites and cross-species amplification is generally lower than expected. This paper details 13 microsatellite loci isolated from the Moluccan Cockatoo and Cuban Amazon and their conservation in other species of parrots.     

Queen palm fruit selection and foraging techniques of squirrels in the Atlantic Forest

Queen palm seeds (Syagrus romanzoffiana) are a vital source of nutrients for Ingram's squirrel (Guerlinguetus ingrami), a common inhabitant of the Atlantic forest biome of Brazil. G. ingrami acts as both a seed predator and disperser of this important palm; however, little is known about how G. ingrami selects or efficiently opens queen palm seeds. The objectives of this study were to: (1) investigate the squirrel's capacity to determine the germinal orifice among the fruit's three pores, which permits more efficient opening of the fruit; (2) assess whether the thickness and size of the fruit endocarp influence the squirrel's palm selection during foraging; and (3) evaluate whether the rodents preferentially select fruits with invertebrate infestations. To answer these questions, we used a series of behavioral experiments measuring patterns of squirrel feeding behavior (fruit opening, fruit selection, and patch selection), controlling for environmental variables. We found that (1) squirrels identify the queen palm germinal pore by its greater relative depth to the other pores; (2) the average size and thickness of fruit does not influence the squirrel's choice of palm for foraging; and (3) squirrels preferentially consume fruits infested by invertebrates. The development of methods for more efficient processing of queen palm fruit highlights the importance of this resource for G. ingrami. In addition, due to its preferential selection of infested fruits, G. ingrami may benefit palms by reducing parasite population loads.     

Diversity of palm uses in the western Amazon

We used palm knowledge to understand the interaction between people and the rainforests and the factors that influence this dynamic process. We interviewed 278 informants in 12 villages in the Pastaza and Madidi areas of the western Amazon basin. Together they used 38 different palm species for 38 different uses in six use-categories (food, construction, utensils, ritual, medicine, commercial). Euterpe precatoria, Iriartea deltoidea, and Oenocarpus bataua were best known and were mentioned as useful by 76–72% of the informants in the 12 villages. There was a great variation in number of uses and in how widespread the uses were and five of the 38 useful palms were mentioned by only one informant. Among the socioeconomic factors analyzed, the residence village influenced the informants’ knowledge of palms and their uses more than any other factor. Length of education, prosperity and for how long the informant had lived in the village were also positively correlated to how many palms and palm uses were known. Gender differences in informants’ knowledge of palm uses were not observed in our data. We suggest, that the high importance of forest products to the livelihood of the villagers, the great variation in the knowledge they possess, and the fact that the differences between villages is so great, are important elements to consider when developing management plans for the sustainable use of the rainforest resources in the western Amazon.     

Extensive consumption of Tabebuia aurea (Manso) Benth. & Hook. (Bignoniaceae) nectar by parrots in a Tecoma savanna in the southern Pantanal (Brazil).

Neotropical parrots forage for various food items such as seeds, fruit pulp, flowers, young leaves, and even arthropods. While foraging, many species wander over large areas that include both open and closed habitats. In this study, I examined parrot foraging activity during a brief synchronous and massive flowering in August 1998 in a tecoma savanna (dominated by Tabebuia aurea) in the southern Pantanal. Six parrot species, ranging from the small Brotogeris chiriri to the large Amazona aestiva, foraged for T. aurea nectar, but Nandayus nenday was by far the major nectar consumer, and the results of each of their visits, like those of the other species, was damage of a substantial proportion of the existing flower crop. Parrots foraged mostly during the afternoon, when nectar concentration tended to be higher. Nevertheless, compared to bird-pollinated flowers, which produce copious nectar, T. aurea had a smaller mean nectar volume per flower. Hence, presumably the amount of damage wreaked by these parrots resulted from their efforts to obtain part of their daily energy and water requirements. Thus, the synchronous and massive flowering occurring in such a brief period in the dry season may be related to, among other factors, the necessity of satiating predators such as parrots, which are still abundant in the Pantanal.     

Characterization of microsatellite loci in the Blue‐and‐gold Macaw,Ara ararauna (Psittaciformes: Aves)

Microsatellite loci were isolated from the Blue‐and‐gold Macaw (Ara ararauna), a Neotropical parrot, from a GT_n and CT_n enriched genomic library. Six loci were characterized varying from one to 11 alleles per locus. Five loci exhibited greater than 50% heterozygosity within the 49 individuals genotyped. Furthermore, the primers also amplified the DNA from two additional genera of Neotropical parrots, indicating the potential utility of these markers for population‐level studies and conservation efforts of Neotropical parrots.     

A comparison of sex steroid hormone excretion and metabolism by psittacine species

The metabolism, excretory rates, and excretory patterns of carbon 14 (¹⁴C) radiolabeled estradiol (E₂) and testosterone (T) were studied in female budgerigars (Melopsittacus undulatus) and orange‐winged Amazon parrots (Amazona amazonica). Radiolabeled E₂ and T were injected intramuscularly into six budgerigars and two orange‐winged Amazon parrots. Serial fecal/urine samples were collected for 168 h post‐radiolabel injection. Peak radiolabeled E₂ excretion was observed at 4 h post‐injection, and by 24 h, 93.3 ± 6.3 and 65.9% (range, 59.1–72.7%) of the injected radiolabel was recovered in the fecal/urine matter of budgerigars and orange‐winged Amazon parrots, respectively. Similarly, peak radiolabeled T excretion was observed at 4 h post‐injection with 92.7± 3.6 and 66.2% (range, 57.5–75.2%) of the injected radiolabel recovered in the fecal/urine matter by 24 h in the budgerigars and orange‐winged Amazon parrots, respectively. High‐performance liquid chromatography (HPLC) analysis of the fecal/urine material revealed that both parrot species excreted >80% of the radiolabel in the form of complex steroid conjugates. Immunoreactive E₂ and T metabolites were detected using estrone (E₁) and C‐21/C‐19 conjugate enzyme immunoassays, respectively. Hydrolysis of the E₂ metabolites and HPLC analysis of the ether extracts revealed that E₂ and E₁ were the major steroid moieties. Hydrolysis of the T metabolites and HPLC analysis of the ether extracts revealed two and three major unconjugated peaks for the budgerigars and the orange‐winged Amazon parrots, respectively. Zoo Biol 18:247–260, 1999. © 1999 Wiley‐Liss, Inc.     

Invertebrate and Vertebrate Seed Predation in the Amazonian Palm Attalea maripa1

Spatial‐scale‐dependent patterns of Attalea maripa seed predation and removal by the beetle Pachymerus cardo and vertebrates were examined in a palm patch and its adjacent forest in southeastern Amazon. Seed survivorship depended on the distance from the palm patch, but was unrelated to distance from individual fruiting palms.     

Geographic variation in the vocalizations of Australian palm cockatoos (Probosciger aterrimus)*

Vocal dialects have been well studied in songbirds, but there have been fewer examples from parrots. The Australian population of palm cockatoos (Probosciger aterrimus aterrimus) from Cape York Peninsula in far north Queensland has an unusually large vocal repertoire for a parrot. Most calls are made during their unique display ritual, which also includes a variety of postures, gestures and the use of a manufactured sound tool. Here, we quantify the geographic structural variation of contact calls within and between six major populations of palm cockatoos in Australia, as well as the extent to which frequently given call types are shared. We found that palm cockatoos from the east coast (Iron Range National Park) possess unique contact calls and have fewer call types in common with other locations. This may have resulted from their long-term isolation in rainforest habitat refugia. Such variety in vocal traits presents a rare opportunity to investigate the evolutionary forces creating behavioural diversity in wild parrots. This is also a step towards assessing links between behavioural variation and population connectivity, which is important information for determining the conservation status of palm cockatoos.     

Energetics of New Zealand's temperate parrots

Abstract

As in most parrots (Order: Psittaciformes) studied, New Zealand species have, independent of body mass, high basal rates of metabolism, low thermal conductances, and precisely regulated body temperatures. An analysis of covariance showed that basal rate in parrots correlates with body mass and thermal climate; temperate species have basal rates that are 21% higher than those of tropical species; and New Zealand's parrot have basal rates 32% higher than expected from body mass. Present information suggests that basal rate in parrots appears to be correlated neither with water availability in the environment nor with food habits. High basal rates in parrots are associated with large pectoral muscle masses. The very small pectoral muscle masses of the kakapo, Strigops habroptilus, predict that this flightless parrot has a low basal rate.     

Disruption of recruitment in two endemic palms on Lord Howe Island by invasive rats

Invasive species may have negative impacts on many narrow range endemics and species restricted to oceanic islands. Predicting recent impacts of invasive species on long-lived trees is difficult because the presence of adult plants may mask population changes. We examined the impact of introduced black rats (Rattus rattus) on two palm species restricted to cloud forests and endemic to Lord Howe Island, a small oceanic island in the southern Pacific. We combined estimates of the standing size distribution of these palms with the proximal impacts of rats on fruit survival in areas baited to control rats and in unbaited areas. The size distribution of palms with trunks was comparable across baited and unbaited sites. Small juvenile palms lacking a trunk (<50 cm tall) were abundant in baited areas, but rare in unbaited sites for Lepidorrhachis mooreana, and rare or absent in 3 out of 4 unbaited Hedyscepe canterburyana sites. All ripe fruits were lost to rats in the small fruited L. mooreana. Fruit removal was widespread but less (20–54%) in H. canterburyana. Both palms showed evidence of a reduced capacity to maintain a juvenile bank of palms through regular recruitment as a consequence of over 90 years of rat impact. This will limit the ability of these species to take advantage of episodic canopy gaps. Baiting for rat control reduced fruit losses and resulted in the re-establishment of a juvenile palm bank. Conservation of both endemic palms necessitates control (or eradication) of rat populations on the unique cloud forest summits of the island.     

Cross-species amplification and optimization of microsatellite markers for use in six Neotropical parrots

Short amplicon primers were redesigned for 17 microsatellite loci developed in St. Vincent's Amazon and six loci developed in blue-and-yellow macaw and tested using six species of Neotropical parrot. Polymorphism was observed at 12 loci in blue-and-yellow macaw, 10 in red-and-green macaw, 11 in scarlet macaw, 10 in chestnut-fronted macaw, 11 in red-bellied macaw and 16 in mealy parrot. Number of alleles per locus ranged from two to 23 and expected heterozygosity ranged from 0.05 to 0.95. The resulting multiplexed loci will be useful in evaluating genetic diversity, genetic structure and mating system in Neotropical parrots.     

Micro‐evolutionary diversification among Indian Ocean parrots: temporal and spatial changes in phylogenetic diversity as a consequence of extinction and invasion

Almost 90% of global bird extinctions have occurred on islands. The loss of endemic species from island systems can dramatically alter evolutionary trajectories of insular species biodiversity, resulting in a loss of evolutionary diversity important for species adaptation to changing environments. The Western Indian Ocean islands have been the scene of evolution for a large number of endemic parrots. Since their discovery in the 16th century, many of these parrots have become extinct or have declined in numbers. Alongside the extinction of species, a number of the Indian Ocean islands have experienced colonization by highly invasive parrots, such as the Ring‐necked Parakeet Psittacula krameri. Such extinctions and invasions can, on an evolutionary timescale, drive changes in species composition, genetic diversity and turnover in phylogenetic diversity, all of which can have important impacts on species potential for adaptation to changing environmental and climatic conditions. Using mtDNA cytochrome b data, we resolve the taxonomic placement of three extinct Indian Ocean parrots: the Rodrigues Psittacula exsul, Seychelles Psittacula wardi and Reunion Parakeets Psittacula eques. This case study quantifies how the extinction of these species has resulted in lost historical endemic phylogenetic diversity and reduced levels of species richness, and illustrates how it is being replaced by non‐endemic invasive forms such as the Ring‐necked Parakeet. Finally, we use our phylogenetic framework to identify and recommend a number of phylogenetically appropriate ecological replacements for the extinct parrots. Such replacements may be introduced once invasive forms have been cleared, to rejuvenate ecosystem function and restore lost phylogenetic diversity.