Imitation of cultural practices is ubiquitous in humans and often involves faithful copying of intransitive (i.e., non-object directed) gestures and societal norms which play a crucial role in human cumulative cultural evolution. Apart from learning these directly from a tutor, humans often learn passively as third-party observers from the interactions of two or more individuals. Whether third-party imitation has evolved outside humans remains unknown. In the current study, we investigated whether undomesticated blue-throated macaws (Ara glaucogularis) could imitate in a third-party setting. A naïve test group (N = 6) passively observed a conspecific demonstrator performing rare intransitive actions in response to specific human gestural commands. Directly afterwards, the observer received the same gestural commands and performance-contingent rewards. An equally naïve control group (N = 5) was tested correspondingly, in the absence of third-party demonstrations. The test group learned more target actions (mean = 4.16 versus mean = 2.2) in response to the specific commands, significantly faster and performed them more accurately than the control group. The test group also spontaneously imitated some of the actions even before they received any gestural commands or rewards. Our findings show that third-party imitation, even for intransitive actions, exists outside humans, allowing for rapid adaption to group specific behaviours and possibly cultural conventions in parrots.Supplementary InformationThe online version contains supplementary material available at 10.1038/s41598-025-11665-9.

Many birds, including macaws, are highly visual animals able to detect a wide band of light wavelengths ranging into ultraviolet A, but in captivity, full-spectrum lighting is not universally employed. Where purpose-made bird lighting is used, this is typically made with the provision of ultraviolet B radiation and vitamin D3 synthesis in mind. Limited research in this field suggests behavioural and physiological benefits of broad-spectrum lighting provision, but more work is needed to broaden the taxonomic scope and to investigate its impacts on understudied areas of husbandry, including behavioural management. We compared the duration of time a bonded pair of blue-throated macaws at ZSL London Zoo opted to remain in an inside den after being recalled from an outdoors flight aviary, with and without the presence of artificial lighting in the form of High Output T5 Fluorescent lamps, which are rich in UVA and UVB wavelengths as well as those visible to humans. We hypothesized that the birds would remain inside for longer when T5 lighting was on, as they would be more visually comfortable. Using randomization analyses, we show that, over 54 trials split between winter and spring, the mean duration spent inside after recall increased from 81.04 to 515.13 s with the presence of the lighting unit, which was highly statistically significant. Our results are likely to be explained by much higher visibility of indoor surroundings creating a more hospitable indoor environment for the birds and will have implications for captive macaw management.

Following a request from the European Commission, the EFSA 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 Allura Red AC for small non‐food‐producing mammals and ornamental birds when used as an additive that add or restore colour in feedingstuffs. The use of Allura Red AC up to the proposed conditions of use of 500 mg/kg complete feed is considered safe for guinea pig, chinchilla, degu, hamster, gerbil and chipmunk. The following maximum safe levels (mg/kg complete feed) apply to the following species: ferrets 99, rabbits 123, canaries, budgerigars, mynah and toucans 45, lovebirds 51, cockatiels 79, cockatoos 115, amazons 145, parrots 147, yellow breast macaw 150, blue‐throated macaw 173 and hyacinth macaw 214. The maximum safe level of Allura Red AC for other small non‐food‐producing mammal is 99 mg/kg feed and for other ornamental birds is 45 mg/kg feed. Inhalation exposure of Allura Red is regarded as hazardous. In the absence of data, the Panel cannot conclude on the potential of Allura Red to be a skin/eye irritant or a skin sensitiser. The FEEDAP Panel cannot conclude on the efficacy of the additive.

The Blue-throated Macaw (Ara glaucogularis) is a Critically Endangered species endemic to the Llanos de Moxos ecosystem of Beni, Bolivia. To aid conservation of the northwestern population that utilizes the Barba Azul Nature Reserve during the non-breeding season, we set out to learn the sites where these birds breed using satellite telemetry. We describe preliminary tests conducted on captive birds (at Loro Parque Foundation, Tenerife, Spain) that resulted in choosing Geotrak Parrot Collars, a metal, battery-operated unit that provides data through the Argos satellite system. In September 2019, we tagged three birds in Barba Azul with Geotrak collars, and received migration data for two birds, until battery depletion in November and December 2019. Our two migrant birds were tracked leaving Barba Azul on the same date (27 September), but departed in divergent directions (approximately 90 degrees in separation). They settled in two sites approximately 50–100 km from Barba Azul. Some details of the work are restricted out of conservation concern as the species still faces poaching pressures. Knowing their likely breeding grounds, reserve managers conducted site visits to where the birds were tracked, resulting in the discovery of breeding birds, although no birds still carrying a transmitter were seen then. A single individual still carrying its collar was spotted 13 August 2021 at Barba Azul. The work suggests that the Blue-throated Macaws of Barba Azul use breeding sites that are scattered across the Llanos de Moxos region, although within the recognized boundaries of the northwestern subpopulation. We conclude that the use of satellite collars is a feasible option for research with the species and could provide further conservation insights.

Forgoing immediate satisfaction for higher pay-offs in the future (delayed gratification) could be adaptive in situations that wild animals may encounter. To explain species-differences in self-control, hypotheses based on social complexity, feeding ecology, brain size and metabolic rate have been proposed. To explore these hypotheses in a comparative setting, we tested three macaw species (neotropical parrots)—great green macaws (N = 8), blue-throated macaws (N = 6), blue-headed macaws (N = 6)—and the distantly related African grey parrots (afrotropical parrots; N = 8) in a modified rotating tray task, in which subjects are required to inhibit consuming a constantly available low-quality reward in favour of a high-quality reward that becomes available only after an increasing delay (min. 5 s, max. 60 s). All four species successfully waited for a minimum of 8.3 s ± 11.7 s (group level mean ± SD) with African greys reaching a delay of 29.4 ± 15.2 s, and great green macaws—as best performing macaw species—tolerating delays of 20 s ± 8 s. The best performing African grey individual reached a maximum delay of 50 s, whereas, a great green and a blue-throated macaw tolerated a delay of 30 s max. Females tolerated higher maximum delays than males. Engaging in distraction behaviours enhanced waiting performance across species and all birds were able to anticipate the waiting duration. Our results suggest that both feeding and socio-ecological complexity may be a factor in self-control, but further systematically collected comparative data on self-control of different (parrot) species are required to test the evolutionary hypotheses rigorously.

A key aspect in the conservation of endangered populations is understanding patterns of genetic variation and structure, which can provide managers with critical information to support evidence-based status assessments and management strategies. This is especially important for species with small wild and larger captive populations, as found in many endangered parrots. We used genotypic data to assess genetic variation and structure in wild and captive populations of two endangered parrots, the blue-throated macaw, Ara glaucogularis, of Bolivia, and the thick-billed parrot, Rhynchopsitta pachyrhyncha, of Mexico. In the blue-throated macaw, we found evidence of weak genetic differentiation between wild northern and southern subpopulations, and between wild and captive populations. In the thick-billed parrot we found no signal of differentiation between the Madera and Tutuaca breeding colonies or between wild and captive populations. Similar levels of genetic diversity were detected in the wild and captive populations of both species, with private alleles detected in captivity in both, and in the wild in the thick-billed parrot. We found genetic signatures of a bottleneck in the northern blue-throated macaw subpopulation, but no such signal was identified in any other subpopulation of either species. Our results suggest both species could potentially benefit from reintroduction of genetic variation found in captivity, and emphasize the need for genetic management of captive populations.

Food availability may vary spatially and temporally within an environment. Efficiency in locating alternative food sources using spatial information (e.g., distribution patterns) may vary according to a species’ diet and habitat specialisation. Hypothetically, more generalist species would learn faster than more specialist species due to being more explorative when changes occur. We tested this hypothesis in two closely related macaw species, differing in their degree of diet and habitat specialisation; the more generalist Great Green Macaw and the more specialist Blue-throated Macaw. We examined their spatial pattern learning performance under predictable temporal and spatial change, using a ‘poke box’ that contained hidden food placed within wells. Each week, the rewarded wells formed two patterns (A and B), which were changed on a mid-week schedule. We found that the two patterns varied in their difficulty. We also found that the more generalist Great Green Macaws took fewer trials to learn the easier pattern and made more mean correct responses in the difficult pattern than the more specialist Blue-throated Macaws, thus supporting our hypothesis. The better learning performance of the Great Green Macaws may be explained by more exploration and trading-off accuracy for speed. These results suggest how variation in diet and habitat specialisation may relate to a species’ ability to adapt to spatial variation in food availability.

Behavioural innovations with tool-like objects in non-habitually tool-using species are thought to require complex physical understanding, but the underlying cognitive processes remain poorly understood. A few parrot species are capable of innovating tool-use and borderline tool-use behaviours. We tested this capacity in two species of macaw (Ara ambiguus, n = 9; Ara glaucogularis, n = 8) to investigate if they could solve a problem-solving task through manufacture of a multi-stone construction. Specifically, after having functional experience with a pre-inserted stick tool to push a reward out of a horizontal tube, the subjects were required to insert five stones consecutively from one side to perform the same function as the stick tool with the resulting multi-component construction. One Ara glaucogularis solved the task and innovated the stone construction after the experience with the stick tool. Two more subjects (one of each species) did so after having further functional experience of a single stone pushing a reward out of a shortened tube. These subjects were able to consistently solve the task, but often made errors, for example counter-productive stone insertions from the opposing end, even in some of the successful trials. Conversely, multiple trials without errors also suggested a strong goal direction. Their performance in the follow-up tasks was inconclusive since they sometimes inserted stones into un-baited or blocked ‘dummy tubes’, but this could have been an attention-deficit behaviour as subjects had not encountered these ‘dummy tubes’ before. Overall, the successful subjects’ performance was so erratic that it proved difficult to conclude whether they had functional understanding of their multi-stone constructions.

The parrots (Psittaciformes, Aves) are a group of colorful, intelligent, long-lived birds with a wide range of body sizes and plumage colors and patterns. One third of the parrot species is threatened with extinction due to habitat loss and the pet trade, a larger percentage than any other comparable bird order. We present the complete genome sequences of 22 species of parrots from 14 genera and 3 families: Anodorhynchus hyacinthinus, Ara ararauna, Ara chloropterus, Ara glaucogularis, Ara militaris, Aratinga solstitialis, Aratinga weddellii, Cacatua leadbeateri, Eclectus roratus, Eupsittula pertinax, Guaruba guarouba, Lorius garrulus, Myiopsitta monachus, Nymphicus hollandicus, Pionus senilis, Psittacus erithacus, Psittacus timneh, Psitteuteles goldiei, Pyrrhura frontalis, Pyrrhura griseipectus, Pyrrhura molinae, Pyrrhura perlata. Genomic data can be used to better understand species identity, hybridization, genetic diversity, and identification of animal products possibly derived from endangered species.

SummaryReliable population size estimates are imperative for effective conservation and management of globally threatened birds like the ‘Critically Endangered’ Blue-throated MacawAra glaucogularis.Endemic to one of South America’s largest grassland floodplains, the Llanos de Moxos in northern Bolivia, the species’ global population size is uncertain. The region’s inaccessibility renders the application of traditional methods for obtaining bird population estimates impracticable or cost prohibitive. We developed a simultaneous, multilocality, double-sampling approach combined with quantitative habitat availability analyses to obtain the first rigorous population size estimate for the Blue-throated Macaw. We established 11 survey areas across its three subpopulations that were visited twice by one team in each subpopulation over a 23-day period in the 2015 dry season and obtained additional count data from two roost sites. We classified suitable habitat (palm forest islands) using Landsat 8 images and CLASlite forest monitoring software. We extrapolated the number of macaws detected (conservative estimate of the total number of macaws [CETN], highest single count [HSC]) per 100 ha of suitable habitat in each survey area to the entire area of suitable habitat in all subpopulations combined, corrected for the species’ range occupancy of 34.3%. The total number of Blue-throated Macaws detected by survey (CETN) and roost site counts was 137. Across all survey areas, the number of macaws per 100 ha of suitable habitat was 4.7 for the first and 4.4 for the second period for CETN and 3.2 and 3.4, respectively, for HSC data. Corresponding global population estimates were 426–455 (CETN) and 312–329 (HSC) individuals. Other recent research and anecdotal data support these estimates. Although it would be premature to propose downlisting the species to ‘Endangered’, our findings indicate that it has a larger population and slightly larger range than previously thought, and that the positive effects of conservation actions are now becoming apparent.

Eight species of macaw (Psittacidae: Anodorhynchus, Ara, Primolius) have previously been reported from Paraguay. We discuss all verifiable reports of the species from Paraguay and provide comment on their statuses. Ara chloropterus and Primolius auricollis are both widespread species known from multiple reports across a wide geographic area. We report minor range extensions within Paraguay for both species. Anodorhynchus hyacinthinus is documented as a breeding resident with a restricted distribution in the Upper Paraguay River basin. The first modern records of Ara ararauna and the first record of Diopsittaca nobilis for the country are documented with photographs. We are unable to find any confirmed records of Anodorhynchus glaucus from Paraguay and thus consider it of possible occurrence. Two species are rejected as erroneously cited, Ara militaris and Ara glaucogularis. Consequently six species in four genera are now confirmed to occur in Paraguay.

Abstract Social complexity may select for socio‐cognitive abilities. The “loose string” task has become a comparative benchmark paradigm for investigating cooperative problem‐solving abilities in many species, thus enhancing our understanding of their evolution. It requires two individuals working together to solve a problem, specifically by pulling the two ends of a string simultaneously to move a reward towards them. A dyad's performance therefore depends on the individuals’ ability to coordinate their pulling action. Many species, including corvids and parrots, have been tested in this paradigm, but most appear insensitive to the exact cooperative nature of the task. We tested another parrot species, blue‐throated macaws, to further our understanding of social cognition in psittacids. Five birds were tested with different partners in a dyadic setting. The study included two control conditions examining the cognitive mechanism underlying their seemingly cooperative behaviour. All birds were able to simultaneously pull the strings, but their performance did not drop when they were denied mutual visual access, and they failed to obtain food when they needed to wait for their partner. Moreover, the parrots decreased their latency to pull with increasing experience. These findings suggest that the birds may have applied an associatively learnt rule, or relied on acoustic cues, rather than coordinating their actions with the partner. This may not necessarily prove a lack of understanding the partner's role, given that their failure to wait in the delay control test might be explained by their poor inhibitory control abilities. Relationship quality (i.e. affiliation and food tolerance) did not influence dyadic success. Future studies are needed in order to disentangle macaws’ potentially limited cooperative abilities from their lack of inhibitory control.

Evaluating management strategies in the conservation of the critically endangered Blue-throated Macaw (Ara glaucogularis)

ABSTRACTAn evaluability assessment of a program to save a critically endangered bird helped prepare the Blue-throated Macaw Environmental Education Project for evaluation and program improvement. The evaluability assessment facilitated agreement among key stakeholders on evaluation criteria and intended uses of evaluation information in order to maximize the utility of an evaluation. Developing a logic model and other activities helped stakeholders agree on program objectives, program description, evaluation options, key evaluation questions, and an evaluation plan. Conducting this low-cost exploratory evaluation exercise before an actual program evaluation is a good strategy to improve program implementation quality and to wisely allocate evaluation resources, particularly for programs with limited budget, staff, or expertise.

Six genera of the Arini tribe form morphologically diverse group termed as Macaws. The presence of bare facial area distinguishes them from other members of the tribe. Genera and species of Macaw group differ in pattern of this bare skin as well as in body size. The genus Ara contains eight extant species. Two of them are classified as medium-sized Macaws and other six species are classified as large Macaws. Based on morphological similarities and differences, large Macaw species can be segregated into three pairs according to their plumage coloration.So far, representative mitochondrial genomes were sequenced only for Ara glaucogularis (blue and yellow coloration), Ara macao (predominantly red/scarlet) and Ara militaris (predominantly green) species. Ara ararauna is the second of two Ara species with predominant blue and yellow plumage. Hitherto, close phylogenetic relation of these two species was shown using only ND2 genes or incomplete combined COI/Cytb/ND2 sequences. Therefore, Ara ararauna mitogenome, which was sequenced in this study, will be indispensable to refine the phylogenetic relationships within Macaws group.

Primolius is a genus of midsized Macaws comprising three species. Blue-headed Macaws (Primolius couloni) are native to eastern Peru, extreme western Brazil and north-western Bolivia. In this study, full mitochondrial genome of considered species was sequenced. It is 16,995 bp long and contains 13 protein-coding genes, 2 rRNAs, 22 tRNAs and a control region. Its comparison with published Blue-throated Macaw (Ara glaucogularis) mitogenome revealed their high degree of identity. Primolius couloni mitogenome is the first complete genomic sequence of this genus. It will be indispensable to refine the phylogenetic relationships within the tribe Arini and will enrich the resource of markers for systematic, phylogenetic and population genetic studies.

The Red-bellied Macaw (Orthopsittaca manilata) is a species of the monotypic genus Orthopsittaca. The genus is one of the six genera, which form morphologically diverse group termed as Macaws. Individuals of Orthopsittaca manilata species are found in extremely large Amazonian area of South America. In this study, full mitochondrial genome of considered species was sequenced. It is 16,985 bp long and contains 13 protein-coding genes, 2 rRNAs, 22 tRNAs and a control region. Its comparison with published Blue-throated Macaw (Ara glaucogularis) mitogenome revealed their high degree of identity.Presented Orthopsittaca manilata mitogenome is the first complete genomic sequence of this genus. It will enrich the resource of molecular markers for future examination of evolutionary diversification of Macaws. It will be also indispensable to refine the phylogenetic relationships within the tribe Arini.

Abstract Monitoring of wild populations is central to species conservation and can pose a number of challenges. To identify trends in populations of parrots, monitoring programmes that explicitly take detectability into account are needed. We assessed an occupancy model that explicitly accounted for detectability as a tool for monitoring the large macaws of Bolivia's Beni savannahs: the blue-throated Ara glaucogularis, blue-and-yellow Ara ararauna and red-and-green macaws Ara chloropterus. We also evaluated the joint presence of the three macaw species and estimated their abundance in occupied areas. We modelled occupancy and detection for the three macaw species by combining several site and visit covariates and we described their conditional occupancy. Macaws occupied two thirds of the surveyed area and at least two species occurred together in one third of this area. Probability of detection was 0.48–0.86. For each macaw species, occupancy was affected by the abundance of the other two species, the richness of cavity-nesting species, and the distance to the nearest village. We identified key priority areas for the conservation of these macaws. The flexibility of occupancy methods provides an efficient tool for monitoring macaw occupancy at the landscape level, facilitating prediction of the range of macaw species at a large number of sites, with relatively little effort. This technique could be used in other regions in which the monitoring of threatened parrot populations requires innovative approaches.

Rediscovered in the wild twenty years ago, the breeding biology of wild Blue-throated Macaws remains largely unexplored, yet is essential to its effective conservation and recovery. Here, we analyse reproductive parameters in an intensively managed wild population of Blue-throated Macaws, providing the first data on the breeding biology of this critically endangered species. During the six-year study period, 2007–2012, the number of active breeding pairs either remained constant or decreased, depending on the site, and no new breeding pairs were discovered despite extensive searching. We documented nesting attempts in natural cavities in dead palms or live hardwoods, and artificial nest boxes. Egg-laying was concentrated during the end of dry season and the beginning of the wet season, August through December. Hatching failure was the greatest cause of egg losses. Half of the breeding attempts of Blue-throated Macaws produced at least one fledging, on average two, after a 85 days nestling period. An average of 4.3 nestlings per year fledged from all known wild nests combined. Each pair lost roughly 65% of its initial reproductive investment at each nesting attempt. In most successful nesting attempts of individualized pairs, a new nesting attempt was not detected the following year. All monitored breeding pairs showed high nest site fidelity, reusing hardwood-tree cavities and nest boxes. Our findings will aid conservation efforts by refining current actions and prompting new approaches towards the conservation and recovery of the Blue-throated Macaw.

Blue-throated Macaw (Ara glaucogularis) is Critically Endangered species of parrot endemic to small Bolivian area. In this study, full mitochondrial genome of considered species was sequenced. It is 16,983 bp long and contains 13 protein-coding genes, 2 rRNAs, 22 tRNAs and a control region. It`s comparison with published Ara macao mitogenome revealed their high degree of identity. On the other hand, analysis of both the genome compositions showed incompleteness of Ara macao CYTB gene. Hence, mitogenome of Ara macao species occurred only partial sequence. In consequence, Ara glaucogularis mitogenome is the first complete Macaw sequence, which will be indispensable to refine the phylogenetic relationships within the tribe Arini and will enrich the resource of markers for systematic, phylogenetic and population genetic studies.