EXITING GENETIC NEWS ABOUT THE SCARLET MACAW (ARA MACAO)
An impressive Ara macao cyanopterus adult bird sitting ready to take off from a piece of rock.
Among the large Central and South American Macaws, the Scarlet Macaw (Ara macao) has over the years become my absolute favorite, especially when it comes
to the subspecies, Ara macao cyanopterus (former known as cyanoptera), in English known as the Central American Scarlet Macaw or Northern Scarlet Macaw (among several other designations). When you look at
this subspecies in top condition and with a perfect plumage, its beauty is beyond compare. In my opinion, no other large Macaw species is able to match these charismatic birds due to their absolute beautiful and sharply demarcated, extremely colorful plumage.
Moreover, it is my experience that they are a bit gentler and calmer compared to some of the other large Macaw species.
The Scarlet Macaw was first described by the great Swedish taxonomist and zoologist (etc.), Carl Linnaeus,
back in 1758 as Psittacus macao, later Ara macao, and it was for more than two centuries considered a monotypic species.
However, in 1994 the American scientist - today
Senior Conservation Scientist at American Bird Conservancy, Virginia, US - Mr. David Wiedenfeld, discovered and described Ara macao cyanopterus (from Central America) as a new subspecies. This discovery has been acknowledged by leading
societies for natural science, museums, institutions, etc. David Wiedenfeld conducted his scientific study based on four morphometric traits/measurements (upper mandible length, lower mandible length, longest toe length and wing chord-length) and on plumage
variation among 31 museum specimens of known geographical origin, and his work led to the division into two subspecies. Ranging from southern Mexico to central Nicaragua Ara macao cyanopterus is considered as bigger and more robust,
with a wide band of yellow and general absence of green plumage on the median and secondary wing coverts. Populations distributed from southern Nicaragua to Brazil are classified as Ara macao macao (with the status as nominate form)
and are identified by reduced yellow and more pronounced green coloration in wing feathers, as well as overall smaller size. David Wiedenfeld also observed a steep decline in wing chord-length and intergradation of color variants in southern Nicaragua and
northern Costa Rica, designating this area as a natural hybrid zone between Ara macao cyanopterus and Ara macao macao.
A wild Ara macao cyanopterus sits outside of a rescue center for Macaws in the small town of Mabita, Honduras (Photo: Art Howard / Christi Lowe Productions).
As I wanted to learn more about the characteristics of Ara macao cyanopterus from a scientific perspective, I started a dialogue with Mr. David Wiedenfeld,
who was very helpful in relation to my many questions. He also drew my attention to a - to me completely unknown - new (late 2019) comprehensive scientific study of the Scarlet Macaw (Ara macao) seen from a genetically point
of view. This new - technically very complicated - study made by a research team of scientists (Kari L. Schmidt et al.) from Colombia University, American Museum of Natural History and Montclair State University, has clearly substantiated David Weidenfeld’s
discovery from 1994 of a new subspecies, this time based on genetic material from 100 Scarlet Macaw samples of birds distributed across the majority of the species’ historical range. This is the newest, most comprehensive and detailed genetic scientific
study ever conducted of the Scarlet Macaw. The study was published in 2019 in "IBIS - International Journal of Avian Science", which is run by the British Ornithologists' Union. Founded in 1858, the British Ornithologists' Union is one of
the world’s oldest and most respected ornithological organizations with an international membership stretching across all continents.
Before I share some of the key findings from this new genetic study with other serious
aviculturists for whom aviculture is not only a hobby, but a passion based on an approximate scientific approach, I would like to emphasize our responsibility as serious aviculturists. We as bird keepers - and not at least as breeders - can make a noticeable
difference by purposefully preserve these rare and beautiful birds in human care (“captivity”). As a serious aviculturist it is of greatest importance in details to know the difference between the two subspecies and to keep them apart from each
other and breed them separately. It is only up in recent times, that aviculturists have become really aware of the two different species. Unfortunately, many aviculturists still don’t know the difference, and it has in an unhappy way led to breeding
of numerous hybrids between the two subspecies in human care over the years. If we - the serious aviculturists - don’t act responsible, it will inevitably soon lead to the annihilation of the latest discovered subspecies, Ara macao cyanopterus,
in human care. We must take good care of those pure birds we already have, as we no longer can get birds from nature. And who knows, over time, we aviculturists may also be able to get our Scarlet Macaws “subspecies determined” based on genetic
analysis as we already know from other non-monotypic parrot genera. In addition, the Ara macao cyanopterus has in recent years faced a number of immediate threats in the wild, partly in the form of destruction of its natural habitats
and its nesting opportunities, and partly as a result of persistent capture for use as pets.
Ara macao cyanopterus, a few months old vital bird with a wide band of yellow and general absence of green plumage on the median and secondary wing coverts. If you keep two or more of these young birds together, make sure that they are continuously stimulated and employed. You should therefore give them lots of new fresh branches and twigs (must not be contaminated and not exposed to pesticides), as they at times may find themselves entertained by biting each other in their very long tails. The tip of the tail may, by virtue of the length of the tail, have a tendency to bend downwards.
And now to the new study:
THE PURPOSE OF THE NEW STUDY
Many people - not least aviculturists - have over the years believed that several different kinds of Scarlet Macaws (Ara macao) exist due to differences in size and in the coloration of the plumage.
More - including authors of books on Macaws - have, without having any scientific evidence, made descriptions of the, in their opinion, several kinds of Scarlet Macaws (Ara macao) that exist. Until now, these authors and others haven’t
had the scientific basis to make credible statements about this subject, but it is something that this new study now is changing. Therefore, this new study is extremely interesting for serious aviculturists.
(Ara macao) have the largest geographical distribution of any Neotropical parrots and occur widely across different habitat types (primary a variety of lowland forest habitats) from Mexico to Bolivia and eastern Brazil. According
to BirdLife International the extent of occurrence (breeding/resident) for the species as a whole is estimated to 10.200.000 km2. Therefore, there is good reason to suppose that this species also is characterized by a number of differences in terms of variances.
Hence the new study aimed to investigate the extent of genetic diversification within Ara macao by analyzing mtDNA sequence data (mitochondrial DNA, the part of the DNA producing energy, and it differs from other DNA, since it
is only inherited from the mother like in most multicellular organisms) from present and historical samples of known geographical origin.
that mitochondrial DNA only is maternally inherited, enables genealogical researchers to trace maternal lineages far back in time. Thus, the evolutionary aspect of the Scarlet Macaw throughout the ages has also been the focus of this
Ara macao cyanopterus, a few months old bird. On this bird you can also observe a wide band of yellow and general absence of green plumage on the median and secondary wing coverts; note that the tips of the secondary wing coverts are blue, there are no green feathers at all. According to Mr. David Wiedenfeld who discovered the new subspecies, Ara macao cyanopterus, this is described as having little to no green coloration, whereas Ara macao macao exhibits small to substantial amounts of green plumage, with no empirical means to quantify “little” versus “small”.
The research team used a number of widely recognized and well-proven scientific tools, methods and processes in order to obtain a solid and well-founded genetic analysis material to provide a credible and secure research
Genetic material (samples) from 100 different Scarlet Macaws distributed across the majority of the species’ very wide range was obtained. DNA was also extracted from feather, blood or tissue collected from
wild individuals in Guatemala and Belize.
A partner (University of Idaho, US) provided DNA-extracts from feathers acquired along clay licks in the Tambopata National Reserve, Peru, which is a well-known territory for Macaws,
Amazons and several other parrot species. The remaining tissues were toe-pads taken from study skins located among natural history museums from all over the USA and handled according to formal processes.
it further, it may be mentioned that DNA was amplified (repeatedly copying a piece of DNA) and sequenced at four mitochondrial gene regions. DNA sequencing is the process of determining the nucleic
acid sequence - the order of nucleotides in DNA.
of reconstructing the evolutionary history of related species by grouping them in successively more inclusive sets based on shared ancestry, called phylogenetic inference analysis, was also used. Actually, two different methods of phylogenetic
inference were used to evaluate hierarchical relationships among haplotypes, maximum likelihood (ML) and Bayesian inference (BI). Haplotypes are a group of alleles in an organism that are inherited together from a single parent, and alleles are each of two or more alternative forms of a gene that arise by mutation and are found at the same place on a chromosome.
The research team furthermore used population aggregation analysis to uncover the current hypothesis of intraspecific diversity in Scarlet Macaws.
Corrected genetic distances
were calculated among each aggregated population identified in the study using four different genetic regions (mentioned above) to further investigate patterns of intraspecific genetic variation.
To assess further taxonomic
distinctiveness and evaluate the extent of divergence between Ara macao cyanopterus and Ara macao macao, the research team succeeded with sequencing a complete
mitochondrial genome from each subspecies (genome is a complete set of genes for an organism).
After the study all the uncovered DNA-sequences were deposited in GenBank®,which is an international genetic
sequence database containing an enormous collection of all publicly available DNA sequences worldwide.
During the new study 70 haplotypes were detected among concatenated mtDNA sequences from the 100 Scarlet Macaws sampled across the species’ range.
Phylogenetic reconstruction describes evolutionary relationships in terms of relative newness
of common ancestry and was used on the sequence dataset, which revealed two major monophyletic groups (a term used to describe a group of organisms that are classified in the same species and share a most common recent ancestor), with near identical topologies
recovered for both “ML” and “BI”, cf. above. The haplotypes within Ara macao cyanopterus formed a well-supported clade (a group of organisms believed to comprise all the evolutionary descendants of a common
Five clusters of haplogroups (haplogroup is a group of similar haplotypes that share a common ancestor with certain other characteristics) were identified withinAra macao cyanopterus
(in the new study referred to as Haplo1, Haplo2, Haplo3, Haplo5 and Haplo6 - regarding the prevalence of the individual haplotypes, see the figure below). Haplo2 was found throughout in upper Central America, from Mexico to northeastern Costa Rica, whereas
the remaining four haplogroups for Ara macao cyanopterus exhibited more restricted geographical distributions. Haplo1 and Haplo6 were both detected in Mexico and Guatemala. Haplo3 and Haplo5 demonstrated similar ranges along the Pacific
slope of El Salvador, Honduras and Nicaragua, however, Haplo3 was also recovered on Isla Coiba, Panama. There is little divergence regarding relationships between Ara macao cyanopterus haplotypes, with the exception of Haplo2 appearing
more diverged from the other four Ara macao cyanopterus haplogroups. Each Ara macao cyanopterus haplogroup clustered into a monophyletic group. Evolutionary relationships among the five haplogroups were poorly
The population aggregation analysis showed no diagnostic characters correspond with the hypothesized subspecies border in central Nicaragua mentioned by David Wiedenfeld (1994). Instead, the geographical limit appears to occur further
south in Costa Rica.
Ara macao macao was characterized by only two haplogroups (in the new study referred to as Haplo4 and Haplo7) with separate geographical ranges demarcated by the Andean Mountains.
The Trans-Andean populations in Colombia and lower Central America belong to Haplo7, whereas Haplo4 is distributed throughout the huge Amazon Basin eastwards.
Here you see one of the figures from the new study showing the prevalence of the individual
haplogroups mentioned above:
In compliance with the broadly sampled phylogenetic reconstructions the research team made, the sequencing and analysis of the full mitochondrial genomes of Ara macao cyanopterus and Ara
macao macaoshowed clear separation of the two subspecies. Concerning molecular diversity average genetic distances between the two subspecies equaled 0,8 %.
The new study can therefore
be said to "stand on the shoulders" of the work that other American scientists were doing back in 2013, when they in Texas initially succeeded in sequencing an overall genome for Ara macao.
Ara macao cyanopterus, an adult beautiful breeding pair in a nice big aviary. On one of the birds, you can see that the underside of the wings is red/reddish, which according to some aviculturists makes it a "true" Ara macao cyanopterus, although this is not substantiated by science.
Evaluation of current taxonomy
The primary visual identifying characters
for the Scarlet Macaw subspecies are listed as a generally larger size for Ara macao cyanopterus and the presence of a green band on the secondary wing coverts for Ara macao macao. However, previous examinations
of morphometric data revealed weak differentiation of geographical variants (cf. David Wiedenfeld, 1994). Only wing chord-length demonstrated statistical significance when corrected for sex, significantly more for males. Likewise, the extent of green tips
on secondary wing coverts, an ambiguous and subjective trait, exhibits a pattern of clinal variation (a gradual change in a character or feature across the distributional range of a species or population, usually correlated with an environmental or geographic
transition); Ara macao cyanopterus is described as having little to no green coloration, whereas Ara macao macao exhibits small to substantial amounts of green plumage, with no empirical means to quantify
“little” versus “small”. Relative to morphological data, a clearer picture emerges when we consider molecular genetic data. There is strong divergence between clades and sequence variation
clearly differentiating the two Scarlet Macaw subspecies.
Ranging from Mexico to northern Costa Rica, Ara macao cyanopterus forms a well-supported monophyletic group consisting of
five monophyletic haplogroups, with each pair differentiated by three to eight fixed nucleotide mutations. The new study shows these distinct haplogroups represent a single evolutionary entity.
populations of Ara macao macao also form a monophyletic group, and they show less pronounced sequence divergence but greater geographical structure relative to Ara macao cyanopterus. Two fixed nucleotide
mutations differentiate Cis-Andean from Trans-Andean populations, with an additional fixed nucleotide delineating Ara macao macao populations in southern Costa Rica and Panama. It is traditional to use the term Cis-Andean for
species distributed East of the Andes (Amazonas) and the term Trans-Andean for those distributed West of the Andes (Central America).
Formal taxonomic descriptions include discussion of possible
introgression between Ara macao cyanopterus and Ara macao macao, with the intergradation of color variants occurring simultaneously with a steep decline in wing chord-length across southern Nicaragua and
northern Costa Rica viewed as evidence of a hybrid zone (cf. David Wiedenfeld, 1994). However, the distribution of mtDNA haplotypes observed in this study demonstrates a general pattern of geographical isolation; no Ara macao cyanopterus
haplotypes were detected on mainland Central America south of the Costa Rican cordilleras (a mountain range). The only instance of Ara macao cyanopterus haplotypes within the Ara macao macao distribution
occurs on the large island, Isla Coiba, located near Panama.
This study found that Ara macao cyanopterus and Ara macao macao haplotypes are separated by the central cordilleras
of Costa Rica, a volcanic mountain range varying in elevation from 500 to over 3.800 m and crosscutting the country, with near complete isolation of the Scarlet Macaw’s preferred lowland habitats along the Pacific and Caribbean slopes, suggesting that,
if present, a present hybrid zone would probably be confined to a narrow geographical area.
The observed different patterns of molecular diversity and wide distribution of mitochondrial haplotypes support Ara
macao cyanopterus and Ara macao macao as diagnosably distinct, reciprocally monophyletic and geographically isolated evolutionary units. To be “reciprocally monophyletic” is a term that typically applies
to analyses of gene tree data from two sister species. If you consider two species, A and B, to say that they are reciprocally monophyletic means that all of the haplotypes sampled from species A are more closely related to each other than any haplotype
from species B, and vice versa.
The research team therefore recommended recognizing Ara macao cyanopterus and Ara macao macao as two distinct subspecies, and - very important
- representing two independent conservation units.
The nominate form of Scarlet Macaw (Ara macao macao), here a male from Bronx Zoo, US. On this specimen you clearly see the large areas of the green color in the different wing coverts, and the greatly reduced number of yellow feathers in the same place, giving a somewhat more variegated impression of the bird. Photo from the Internet.
Patterns of diversification
Fluctuations in the distribution of habitats in response to changes in temperature and precipitation are commonly considered to be
key factors governing the historical biogeography of lowland ecosystems across the Central and South America.
Regional differences in the intensity and duration of population fragmentation and associated demographic changes provide a theoretical framework
to explain patterns of evolutionary divergence within Scarlet Macaws. In the case of Ara macao cyanopterus, the subspecies’ range encircles the Central American landmass, geographically constrained by an extensive system of
central highlands and the coastal waters of the Atlantic and Pacific Oceans. The Scarlet Macaws’ preferred humid lowland habitats are heavily dependent on the availability of fresh water, and thus shifts in distributions would closely follow this critical
resource. The genetic signatures recovered among mtDNA haplotypes may reflect the fragmentation of ancestral Ara macao cyanopterus into allopatric refugia (non-overlapping distribution areas during periods of unfavorable conditions).
The presence of several distinct Ara macao cyanopterus haplogroups appears to increase levels of overall sequence variation, as diversity indices decline for the majority of haplogroups when treated as separate units.
different pattern of genetic variation emerges upon examination of Scarlet Macaw populations across lower Central and South America. Ara macao macao is represented by only two haplogroups, differentiated by two fixed nucleotide bases
and geographically separated by the Andean Mountains.
Conservation implications and threats
Common throughout its global distribution through the ages, populations of Scarlet
Macaws have experienced widespread demographic declines and local extinctions due to capture for the pet trade and habitat destruction. Furthermore, anthropogenic pressures have been most severe in Central America, where high human population densities and
associated resource demands put tremendous strain on the region’s ecosystems. This has resulted in a 22 % reduction in primary forest within a 20-year period from 1990 to 2010 (FAO 2010).
The confirmation of genetic diversity reveals a critical
gap between conservation needs and conservation status for these charismatic parrots. At present, Scarlet Macaws are considered a species of “Least Concern”, given global demographic trends, population size and extent of available habitat are above
designated thresholds for threatened status (IUCN 2015). Splitting Scarlet Macaws into two distinct conservation units, however, immediately transforms the conservation status of this group. Approximately 83 % of preferred lowland habitats are located within
the Amazon Basin, along with the majority of the estimated census population of 20.000 - 50.000 individuals for Ara macao (BirdLife International 2011). Deforestation rates are declining throughout South America, dropping from 0,49
to 0,41 % in the past decade (FAO 2010), with human population growth rates falling in concert from 1,17 % to 1,07 % (CEPAL 2013). Therefore, robust populations of Ara macao macao ranging across the Amazonian lowlands may indeed qualify
for “Least Concern” status.
Opposite, the situation faced by Scarlet Macaws in Central America is far more precarious. Annual human population growth rates are currently 1,59 % and human
densities are 4,2-fold greater relative to South America, putting tremendous strain on the region’s natural resources. Forest area in Central America declined by an average of 1,19 % annually between 2000 and 2010, the highest rate reported
by the United Nations (FAO 2010). The loss of important foraging and nesting habitats, coupled with intense nest poaching for the pet trade, have decimated Ara macao cyanopterus with fewer than 4.000 wild Ara macaocyanopterus remaining in isolated forest fragments throughout upper Central America. Recovery of five unique mitochondrial haplogroups within the Ara macao cyanopterus lineage highlights the evolutionary
significance of these populations, further advocating that Ara macao cyanopterus should be up listed to “Vulnerable” status.
Rigorous delineation of conservation
targets is needed
The new study also pointed out that current conservation designations may need to be re-evaluated. The use of the term “subspecies” in taxonomy and conservation biology is considered somewhat controversial given
the arbitrary assessment criteria’s and poor connections to evolutionary history. Therefore, rigorous delineation of conservation targets within Scarlet Macaws is needed to create a solid foundation for conservation assessment, management strategies,
ease of daily handling, etc. The new study has helped to create such clarity, and a clear definition of a subspecies provides a unique conservation target, which hopefully may help to save it in its natural habitats in the long run.
As already stated, it is estimated that fewer than 4.000 Ara macao cyanopterus remain in wildlife in Central America, scattered across areas in smaller isolated forest fragments which makes it very vulnerable. In
comparison, BirdLife International estimates that there are 4.300 Hyacinth Macaws (Anodorhynchus hyacinthinus) left in the wild with a much larger range of 2.850.000 km2, and it is categorized as “Vulnerable”.
is my assessment that if Ara macao cyanopterus in the future can become an independent wild conservation subject, the chances for protecting these wonderful birds in the wild will be much better. However, BirdLife International still
only operates with the Scarlet Macaw (Ara macao) on species-level. As the Scarlet Macaw (Ara macao) in general is widely distributed and thought to be relatively tolerant of degraded habitat, the species
is considered of “Least concern” by the IUCN (2016) and BirdLife International though it also is listed under Appendix I of CITES.
In this area, the United States is significantly further ahead; in 2019 Scarlet Macaws (Ara
macao) received protection under the United States Endangered Species Act (ESA). The northern subspecies of the Scarlet Macaw (Ara macao cyanopterus)
has thus been listed as endangered and a distinct population segment
(DPS) of the southern subspecies (Ara macao macao) as threatened. The southern subspecies (and strangely enough subspecies crosses as well) of the Scarlet Macaw are also added to an existing special rule for parrots under section
4(d) of the ESA. This continues to provide needed protections while allowing for interstate commerce and the import and export of certain captive-bred birds provided the requirements of the Wild
Bird Conservation Act and the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) are met. So, the Scarlet Macaw is now protected
by both this important U.S. conservation law and by the international conservation agreement.
SOME CONCLUSIONS ON THE STUDY
This is also a Scarlet Macaw with a nice yellow wing band, but notice the relatively few green feathers. According to the scientific description of Ara macao cyanopterus, a certain amount of green is acceptable. It is therefore important that you also use the well-known morphological visual features that characterize this subspecies to be more certain that it is an Ara macao cyanopterus and not a hybrid bird
In 1994 the scientist David Wiedenfeld proved the existence of two different subspecies of Ara macao through studies of purely morphological features and
color variation in the plumage. The new scientific study (2019) from Columbia University etc. has through genetic analysis confirmed the existence of these two - and only two – subspecies, and this new study has even revealed strong phylogeographical
divergences within the Scarlet Macaw (Ara macao).
Together with phylogenetic reconstruction and population aggregation analysis the new study has thus confirmed the existence of two distinct phylogeographical
groups, with a high degree of intraspecific genetic structure and surprisingly no evidence of a putative hybrid zone as David Wiedenfeld’s study from 1994 had otherwise assumed. Instead,
it is rather considered to be an actual physical segregation between the two subspecies that occur further south in Costa Rica
Up until now the two subspecies have been weakly differentiated, but this has changed with the
new study. Through the new study the whole mitochondrial genome sequence has been uncovered and has confirmed a substantial divergence between the Ara macao macao- and the Ara macao cyanopterus-lineages.
The huge geographical range of the Scarlet Macaw’s preferred habitats has thus undergone significant changes across evolutionary time, caused by major geological events and climatic fluctuations through the ages. The results
show a separation of Ara macao into two distinct evolutionary entities and highlight a dissimilar distribution of intraspecific diversity, especially in relation to Ara macao cyanopterus caused by phylogeographically
conditioned fragmented presence in its much smaller habitat areas. The Ara macao cyanopterus thus lives in several isolated populations. Although each population now may be large enough to avoid genetic inbreeding problems, because
the populations are small and isolated, their long-term survival unfortunately were already assessed unlikely by David Wiedenfeld (1994).
Ara macao macao was characterized by only two haplogroups
with separate huge geographical ranges divided by the Andean Mountains. On the other hand, Ara macao cyanopterus consists of 5 different haplotypes according to the new study, which means a significant differentiation
in its genotype, which - in my humble opinion - most likely will be visible through higher differentiation in its phenotype, typically shown as differences in the coloration of the plumage.
With these two scientific studies
of the Scarlet Macaw (Ara macao), serious aviculturists now have much better possibilities to distinguish the two subspecies from each other and thus breed with each subspecies separately. However, the scientific work so far is not
quite enough to in practice always be able to distinguish the two subspecies (without genetic tests). Over time, knowledge of other visual characteristics per subspecies has been built up among particularly interested aviculturists. Below is a brief overview
of some of these non-scientific characteristics, or rather perceptions, although not all of them are valid in my opinion.
VIEWS ON ARA MACAO CYANOPTERUS AMONG SOME AVICULTURISTS
Ara macao cyanopterus, a few months old birds. In Denmark, the wind blows very often and much, so much so that the birds - when sitting in outdoor aviaries - almost push themselves down along the branch on which they sit with their beaks pointing towards the wind, as otherwise they will not be able to control their very long tails, which precisely characterizes this subspecies.
Over recent years some aviculturists, that have shown interest for - or even are keeping/breeding - Ara macao cyanopterus, have either heard about or have
developed many perceptions of how this subspecies should look like, their size, tail length and width, etc.
Some of these perceptions are listed in the following and they affect
both morphological (physical) features and color differences in the plumage, that should characterize this subspecies. Be aware that these perceptions are not supported by scientific evidence, but typically solely rest on “reports and
rumors” among aviculturists, etc. Nevertheless, some of them might provide a more complete picture of how a “true” Ara macao cyanopterus should look like, when one combines it with the scientific phenotypic features
mentioned above, so it hopefully better can be visually distinguished from the nominate form (Ara macao macao).
a) Morphological (physical) features:
The perception that the tail feathers on the Ara macao cyanopterus should be of a certain length and width are not supported
by scientific evidence in the two studies mentioned in this article, but of course this subspecies according to David Wiedenfeld should be more robust (a single very big captive bird, for example, has weighed up to nearly 1.600 grams) and longer than Ara
macao macao. The tail appears disproportionately long and among aviculturists there seems to be a common opinion that the tail length on Ara macao cyanopterus most often should be 60 - 70 cm or more. The width of the
longest tail feathers on males can measure up to 5 - 6 cm, on some occasions even wider, in certain areas of the feather.
Another significant difference seems to be that the whole tail of the Ara macao cyanopterus
apparently appears to be of a certain width throughout its length, whereas the tail of the nominate form, the Ara macao macao, appears to be more wedge-shaped as e.g., seen among a number of South American parakeets for example the
Even the physical structure of the tail of an Ara macao cyanopterus is also said to be very different compared to the tail on an Ara macao
macao. It is said that the middle part of the tail on the Ara macao cyanopterus consist of two long center tail feathers, and the tip may, by virtue of the length of the tail, have a tendency to bend downwards. In contrast,
the middle part of the tail on the Ara macao macao should consist of a single centered tail feather.
Some aviculturists believe, that the Ara macao cyanopterus coming
from Nicaragua - in general or at least in some cases - seems larger than other specimens coming from other parts of the range of Ara macao cyanopterus. Several samples of Scarlet Macaws from Nicaragua were used in both scientifical
studies (1994 and 2019) and no notes have been made to substantiate this perception by scientific evidence. Among a large population of birds there will always be a variation in size, coloration, etc., and some individuals will seem bigger and more colorful
than others. This is far from always due to genetic conditions, but instead this may be due to the fact that the birds have had different living conditions, for example differences in the nutritional value of the feeding during the upbringing, the extent and
nutritional value of the feeding in daily life, the birds you compare have different ages, sex, different access to nesting facilities, etc. However, local populations of large types of certain other Macaw species are not unknown: For many years it has been
common knowledge among aviculturists, that specimens of both the Blue and Yellow Macaw/Blue and Gold Macaw (Ara ararauna) and the Red and Green Macaw/Green-winged Macaw (Ara chloropterus) that come from Bolivia
appear to be significantly larger than similar birds that come from other parts of the large range of these two species. On the other hand, the Bolivian specimens of these two species are not significantly different in the coloration of the plumage apart from
minor exceptions. Both of these species are monotypic species, as science does not recognize any subspecies of either Ara ararauna or Ara chloropterus. In these cases, it is a matter of species variance.
Finally, a few aviculturists claim that certain Scarlet Macaws that come from Bolivia in some physical body areas also show larger size.
A huge and adult Ara macao cyanopterus, an impressive, robust male bird with a very long and wide tail that characterizes an Ara macao cyanopterus.
b) Color differences in the plumage:
As I write - to my knowledge - there is no scientific evidence, that the underside of the wings of Ara macao cyanopterus only should be red/reddish, while the underside of the wings of Ara macao macao
should be yellow/yellowish. The study that David Wiedenfeld completed in 1994 does not shed light on such a difference. When evaluating the significance of this alleged characteristic you should - according to my own experience - take into account what age
the bird is. Next to this, it is of utmost importance that two (or more) different birds are compared under identical lighting conditions. Depending on whether you observe the underside of wings in day light (including flash light) or in clouded weather reddish
colored underside of the wings can seem yellowish in bright, sunny weather and yet perceived as red/reddish in clouded weather.
When you observe - or rather examine - a Scarlet Macaw the age of the bird is also important to know, because the colors
of this bird's plumage to a certain extent may change over time, especially when it comes to birds in human care as their life span often are much longer than birds living in the wild. Not least in the birds' first years of life, there may be visible changes
in the coloration of the plumage. Conversely, large color differences can also occur among very old individuals, where the yellow feather areas in the wings of some specimens may vanish and instead replaced by red feathers.
The large color differences that often can be observed in the plumage of Scarlet Macaws have over the years got aviculturists (not least in the US) to group these birds into 3 or even 4 different types depending on their place of origin. Regardless
of such groupings, we now know for sure that science has proven that two - and only two - genetically very different subspecies
of Scarlet Macaws exist. Nevertheless, in nature we can also observe differences in the looks of Scarlet Macaws sitting together in a flock in the same place, but these must be considered as natural varieties as we know it from many other kinds of animals.
However, it should not stop me from briefly mentioning one of these non-scientific groupings based on the birds' different coloration (phenotype) place of origin in nature. Here is an American example of a non-scientific grouping of Scarlet Macaws into three different types:
It's unfortunately not the clearest photo one could have wished for however, on these few months old young birds which are bathing in the rain, you can see that the underside of the wings is red/reddish, as some aviculturists believe they must be in order to be a "true" Ara macao cyanopterus, but there is currently no scientific evidence for this view.
The Mexican Scarlet Macaw-type:
They are quite colorful and are characterized
by a narrow yellow band of feathers on the wing nestled between the red feathers on the shoulder and the blue flight feathers, generally only two rows wide (the edges of the yellow wing coverts being colored with shades of blue to green). These are not huge
birds overall; a collection of Mexican Scarlet Macaws has shown an average weight of 908 grams. Their range has been considerably reduced in Mexico due to deforestation for agriculture, now limited to where there still are jungles. At one point, they ranged
all the way up the east coast of Mexico to just before the northern deserts. Most of the Mexican type still living are in small local forested areas of southern Mexico.
The Central American Scarlet
The Central American Scarlets Macaws come from lower Central America (Nicaragua, Costa Rica, parts of Honduras and Panama, and occasionally as far north as Guatemala) and are considered
the largest and most spectacular Scarlet Macaws to be seen. Some people have been bragging of tail feathers up to 90 cm. Central American Scarlets range in weight from 1.190 to 1.560 grams for males and 1.048 to 1.162 grams for females. These large birds have
the widest band of yellow feathers that when combined with the long tail feathers identify them as Central American Scarlets. The yellow wing band is three to four feathers wide and edged by greenish-blue tips, but only a slight amount of the tip of the yellow
feather is covered by this greenish-blue color.
The South American Scarlet Macaw-type:
South American Scarlet Macaws are generally not as
large as Central American Scarlet Macaws and have a narrower band of yellow feathers, very similar to the Mexican Scarlet Macaw-type, that is bordered in green and - in some specimens - also blue. The weight is average 1.219 grams for males and 1.162 grams
for females. Their range in nature have been reduced significantly, until the last vestiges are in the Amazon Basin, at elevations under 400 m. The marking that differentiates this type from the other two is a large turquoise-green area, commonly the size
of a tablespoon, found on the nape of the neck.
To determine whether a certain Scarlet Macaw belongs to one or the other of the two subspecies can be difficult, and the many
hybrid bred birds among aviculturists make it even more difficult. A Scarlet Macaw with completely yellow median and secondary wing coverts is not always an Ara macao cyanopterus. You must - as described in this article - also look
into other criteria’s as well, mainly morphological traits such as the size of the bird, the size of the beak and especially the length of the wing chords and of course the length and width of the tail, and not to forget - the weight. Therefore, strive
to keep the two subspecies separated from each other when breeding Scarlet Macaws.
A general description of the Scarlet Macaw can be found in several serious non-fiction books such as the magnificent, “Parrots of the
World” (1st edition 1973, or later editions), by the excellent Australian ornithologist, Joseph M. Forshaw, to which reference is made.
Written / Updated: 31.01.2021/31.01.2021
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About the Blue-naped Parrot (Tanygnathus lucionensis)
A story about determining the right subspecies
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My first encounter with the Tanygnathus genus
Already when I was a teenager back at the beginning of the 1970's, you could occasionally see some very special parrot species at one of the large bird traders in Copenhagen (Denmark), who were known for continuously importing many parrots from the various parrot continents. They were smaller medium to medium sized green parrots with a beautiful mosaic pattern on their wings, which is caused by the different colored wing coverts are being broadly marked with orange-yellow edges (in most of the subspecies) and with an impressive, disproportionately large red beak compared to the rest of the bird. These birds later turned out to be Tanygnathus lucionensis and Tanygnathus megalorynchos, and I remember that on the few occasions I saw them, it was usually just a single bird in the cage tucked away over in a corner of the shop premises, possibly because they were mistakenly included in a shipment from Asia along with a number of other far more popular parrot species.
At this time, nobody really knew anything about these birds, and at the same time they did not seem to have much interest among aviculturists. This has since changed quite a bit because nowadays many aviculturists are fascinated by these parrots.
But what is it that makes parrots from the Tanygnathus genus so fascinating?
Tanygnathus lucionensis lucionensis: The scientific Latin species name "lucionensis" means coming from Luzon, the largest island in the Philippines. This is the nominate subspecies coming from the islands of Luzon and Mindore in the northern parts of the Philippines. The beautiful photo clearly shows the upper side of the nominate subspecies which is characterized by it is having a blue/bluish mantle, back, lower back and upper rump. Photo: Forest Botial-Jarvis-Bataan, Philippines-Macaulay Library ML560592651.
Brief introduction to the whole Tanygnathus genus
Before we delve into the substance and take a closer look at the main focus of this article, namely the determination of the Tanygnathus lucionensis subspecies, here is an overall and very short introduction to the entire Tanygnathus genus:
The Tanygnathus genus represents a group of parrots with predominantly green plumage that are smaller medium to medium sized birds with large - in several instances - very large, heavy mainly red beak (hence the Latin scientific designation, “Tanygnathus”, that means extended, long and beak/jaw, is well-deserved) and a proportionately short, wedge-shaped rounded tail, which is somewhat stepped and is shorter than their wings, giving them - according to the world-renowned Australian ornithologist Joseph M. Forshaw - a “top-heavy” look. There is no prominent notch in the upper mandible and the cere is naked. Sexual dimorphism is slight or absent, and young birds generally appear duller than adults.
The premise of this article is the world's leading, current nomenclature/taxonomy, “THE HOWARD AND MOORE COMPLETE CHECKLIST OF THE BIRDS OF THE WORLD”, version 4 from 2013 and newest version 4.1 from August 2018, ”Errata and Corrigenda to Volume 1”, (used by the world's leading scientists, researchers, ornithologists, curators, etc.) mentions the following species and subspecies belonging to the Tanygnathus genus:
Status in the wild according to BirdLife International: “Vulnerable”.
The members of the Tanygnatus genus are closely related to the Eclectus Parrots (the Eclectus genus). However, the most Tanygnathus species are much smaller and are less heavily built than the Eclectus Parrots and thus more elegant. Just as you see with members of the Eclectus genus, Tanygnathus females are also known to be quite aggressive towards males, but in fact some Tanygnathus males can also be aggressive towards females. It can even become so violently that the female's toes can be bitten off. As with the Eclectus Parrots, there is only limited interaction between the sexes in daily life and mutual preening is not something that I yet have experienced in the Tanygnathus species that I have kept for some years.
Genetic evidence has also supported close relationship to the Psittacula genus as it looked like up until 2019. For example, there are many similarities between Tanygnathus megalorhynchus and Psittacula eupatria magnirostris (nowadays Palaeornis eupatria magnirostris, in english Andaman Alexandrine Parakeet); if one abstracts from the fact that the former has a short tail and the latter has a long tail, just try to compare the enormous and amazing beaks of both birds.
This genus is native to Southeast Asia and Melanesia. The overall distribution area is mainly the Philippines, New Guinea, the Moluccas and Sulawesi (the world's 11th largest island, also called Celebes).
In nature, their food mainly consists of fruits (e.g., mangoes), seeds, berries, nuts and insects. Sometimes their food also consists of crops e.g., corn.
What particularly fascinates me about this genus is their disproportionately large beak, which in the vast majority of cases is large and red and thus very prominent, you can't help but focus precisely on it when you look at the bird. It cannot be said that the members of the genus Tanygnathus are brightly colored, but most species/subspecies have an incredibly beautiful mosaic pattern on their wings, which is caused by the different colored wing coverts are being broadly marked with orange-yellow edges (in most of the subspecies). Although some of the birds may appear a little compact, they are very agile and move incredibly elegantly in the branches of the trees.
Kept in human care, these birds appear very shy at first, but gradually they become calmer and, in some cases, they can become completely comfortable with human contact.
None of the four different Tanygnatus species have ever been common within the aviculture, which may be surprising, as they are rather quiet and low-voiced birds that you can become familiar with over time. Having said that, Tanygnathus lucionensis is the "most frequently" occurring species among aviculturists, whereas Tanygnatus megalorynchos and Tanygnathus sumatranus are seen even less often. Very remarkably, Tanygnathus lucionensis has the reverse status in nature where it has become a fragile species with a status of being "Near Threatened". BirdLife International has estimated that there are only 1,500 - 7,000 individuals left in the wild of this species with associated subspecies. Besides, the population development is also decreasing mainly due to logging of trees in its habitats. Neither Tanygnatus megalorynchos nor Tanygnathus sumatranus is nearly threatened in the wild, fortunately they are of the least concern. It is therefore very paradoxical that Tanygnathus lucionensis is the most widespread species among aviculturists in Europe.
Tanygnathus megalorhynchos sumbensis: This photo shows a subspecies of the biggest species belonging to the Tanygnathus genus, the Great-billed Parrot (Tanygnatus megalorynchos) which also have the largest and most impressive beak of these species. Tanygnatus megalorhynchos sumbensis, comes from the island of Sumba (Lesser Sundas). Photo: Mehd Halaouate.
Lack of reliable knowledge about the Tanygnathus genus
One thing is to have a scientifically worked-out taxonomy that lists and divides the Tanygnathus genus into species and subspecies distributed by geographical distribution areas, another thing is to find reliable sources (field studies, literature, photos, etc.) that describe all the members of this genus in detail and accurately. I have many parrot books, encyclopedias, etc., but I have never managed to find a book with accompanying photos that in a credible manner in detail describe all the various Tanygnathus species with its associated subspecies. At best, this genus is all too often referred to be only peripherally described in these books. Moreover, a number of these descriptions are only superficial, and it is of no use if you have to determine the subspecies of a species that has a number of very comparable subspecies (cf. later). The different parrot books contain almost no photos where one can really compare all the subspecies and see the differences between these. Up until now I haven't been able to find a book or another source of sufficient credibility and accuracy.
With this article, I will try to contribute to creating a little more clarification in relation to some of Tanygnathus lucionensis' subspecies, and that is the focus for the rest of this article. At the same time, it must be emphasized that the present article predominantly deals with taxonomic and morphological issues in order to try to determinate the right subspecies of the Tanygnathus lucionensis that I keep, to which comes some general information about the species in the wild. However, the article does not deal with topics such as keeping, feeding and breeding of Tanygnathus lucionensis in human custody.
A little about Tanygnathus lucionensis in the wild
Very summary, this is a smaller medium sized parrot, around 31 cm in length, primarily green except for a light blue rear crown and nape, pale blue lower back and rump (cf. subspecies later), shoulders with broad orange-yellow edges on black wing coverts, and blackish underwings with green underwing coverts.
As already mentioned, this species is listed as “Near Threatened” by BirdLife International because there are some indications that it has a moderately small, fragmented population, and it may be undergoing a continuing decline due to habitat loss (logging) and trapping. However, little is still known about the population size, population structure and threats to this species.
Tanygnathus lucionensis is present throughout the Philippines, where there are records from at least 45 islands, plus the Talaud Islands, Indonesia, and islands off north-east Borneo belonging to Malaysia (e.g., Kota Kinabalu, which is a very popular photo spot for tourists interested in this species) and it is therefore not a country endemic species.
It was common on most islands in the Philippines a century or less ago, but has suffered declines since on such a scale that it is now rare or extinct on many islands. However, it does survive in small pockets of habitat on the smaller islands, so that its status overall is difficult for BirdLife International to assess. Moreover, it is still fairly numerous in some areas of Palawan and on Tawi-Tawi, and high numbers should be present in a large tract of forest on Talaud. Intensive habitat loss and trapping have made this species scarce on most islands except Mindoro and Palawan, but still a local animal protection organization has raised concerns over the increasing illegal trade of this bird on Palawan.
It also occurs in urban areas such as several national parks within the Philippines: Bataan, Quezon and Minalungaw.
As previously stated, BirdLife International's current estimate of the number of Tanygnathus lucionensis (incl. subspecies) is 1,500 - 7,000 mature individuals, while the population development is described as declining owing to habitat degradation from agricultural expansion and logging pressures to which comes illegal trapping for the cagebird trade. It is a bird of closed and open forest formations, including secondary forests, coconut and banana plantations and mangrove, chiefly in lowland and coastal areas, up to 1,000 m. It is usually found in flocks of up to 12 individuals which roost communally and make regular dawn and dusk flights between feeding and roosting areas. Breeding takes place in a hole in a tree in from April till July.
Tanygnathus lucionensis lucionensis: When you see photos of these birds from the nature, you most often see them sitting in the tall tree crowns where they prefer to be. Although some of the birds may appear a little compact, they are very agile and move incredibly elegantly in the branches of the trees. Photo: Stephan Lorenz-Bataan, Philippines-Macaulay Library ML563667891.
The scientifically recognized subspecies of Tanygnathus lucionensis
“THE HOWARD AND MOORE COMPLETE CHECKLIST OF THE BIRDS OF THE WORLD” (see above) recognizes the following subspecies of Tanygnathus lucionensis:
Note to the above table:
I am fully aware that in the spring of 2023, Arndt-Verlag has issued a new - and final - complete version of the "Namensliste der Papageienarten und Unterarten" (Latin, German and English names). It states the existence of - in relation to Arndt-Verlag's poster with the Tanygnathus genus (“Großschnabelpapageien”) - four "other" Tanygnathus lucionensis subspecies, namely nigrorum, siquijorensis, koikei und paraguenus, without these being further described. These four 4 "other" Tanygnathus lucionensis subspecies are not recognized today by the leading scientific taxonomy, and the deviations that the designation of these subspecies has expressed in its time are today simply considered variance within the framework of the currently recognized subspecies. They are thus assimilated into the current range of subspecies.
So, these four "other" Tanygnathus lucionensis subspecies are ignored here, as the article is based on the world's leading taxonomy, which is based on solid scientific evidence as the basis for the division into species and subspecies (for example, the subspecies, siquijorensis, was in its time solely determined from only a single holotype specimen).
All of the four "other" subspecies mentioned by Arndt-Verlag, koikei and paraguenus, as well as nigrorum and siquijorensis, were all already mentioned as subspecies in the world's first large complete parrot book, called "Papegøjebogen" (in English "The Parrot Book"), which was written by the Dane, J. L Albrecht-Møller, with the assistance of another Dane, Aage V. Nielsen. J. L. Albrecht-Møller was approximately 30 years to write "The Parrot Book", which was completely published in 1973. Unfortunately, the book is only available in Danish and consists of almost 700 densely written pages spread over 3 volumes as well as a completely extra 4th volume filled with color drawings of a large number of species and subspecies of parrots of all kinds.
In fact, it was a third Dane, an internationally highly respected scientist, Finn Salomonsen (zoologist and ornithologist), who first described the subspecies nigrorum and siquijorensi. During an expedition to the Philippines in the years 1951 - 1952, he also did some research on this genus.
As mentioned, none of the above four "other" subspecies have found grace for inclusion in Howard and Moore's taxonomy. Nor does the world's second major scientific "heavyweighter" within bird taxonomy, "The Clements Checklist of Birds of the World", 6th Edition, that was published and released by Cornell University Press in June 2007 take these four "other" subspecies into account.
Tanygnathus lucionensis lucionensis: As a nest these birds use a hole in a tree where they during April - July are breeding. The nest is often found in secondary forest, at forest edges and in plantations at elevations of up to 1000 m. Photo: Bradley Hacker-Bataan, Philippines-Macaulay Library ML214490541.
My first purchases of Tanygnathus lucionensis did not go as hoped
From my first encounter with this genus back in the early 1970's, it was always in the back of my mind that when the right opportunity presented itself, I would like to acquire such birds, but almost a lifetime should pass before thoughts turned into action in autumn 2018. Here I acquired my first pair of Tanygnathus lucionensis from a foreign breeder who had guaranteed me a 100 % perfect and unrelated pair. As he himself only had one breeding pair, he had procured an unrelated female bird via another aviculturist in a neighboring country. The breeder informed me that the birds were of the subspecies Tanygnathus lucionensis talautensis, which, compared to the nominate subspecies, that has a blue/bluish mantle, back, lower back and upper rump, instead has a pure green plumage in the same places. He further stated that it was the subspecies that was found most among aviculturists in Europe. After driving many hundreds of kilometers to collect the birds, it unfortunately turned out that the breeder's own male bird - in contrary to what he had otherwise promised - was not 100 % perfect, as it was basically missing half the length of the tail, which I had not been informed about before my departure from Denmark. Since I had driven so far, and because it was a rather rare bird, I chose, after consultation with my wife, who went along on the trip, to bring both birds home to Denmark, but my disappointment with this seller, who, after all, several times had guaranteed that both birds were 100 % in order, was great. At home in Denmark, the birds were placed in an approximately 6-week long quarantine period and different clinical disease testings were conducted both via PCR and serology, and all test results were negative for ABV/PDD, APV, PBFD, and CPS, etc. Already during the quarantine period, I had observed that the male bird, which unfortunately was hand-reared, had an abnormal behavior. Besides, the fact that it had a habit of plucking its own feathers, it was obviously also mentally disturbed, which resulted in highly unusual behaviour, where it, among other things, sat in the same place around the clock without moving, and it seemed almost terrified of e.g., natural branches, to which was added the fact that it had absolutely no interaction with the female bird. I contacted the seller to return the bird, but he refused to take it back, so as the bird's condition further deteriorated, in consultation with a veterinarian specialized in bird diseases, it was decided to euthanize the bird, so my first attempt at acquiring a pair of this species ended in failure. The female was subsequently sold to another aviculturist who was missing a female bird, as I was of the opinion that it probably would be easier to buy a new pair together.
I had not given up on my dream of keeping this species, but next time the acquisition should be made from a much more serious and experienced breeder of this species. During my search for such a breeder, I sometimes came across other aviculturists who also mentioned that the subspecies Tanygnathus lucionensis talautensis is the kind that is most widespread among European aviculturists.
In autumn 2021 - after being registered with a new breeder for nearly 1 year in order to be allowed to buy a pair - I ended up buying a new pair. This breeder sold his birds just as being Tanygnathus lucionensis i.e., without being subspecies determined, but it was clear that even these newly acquired birds did not have the features known for the nominate subspecies in the form of a blue/bluish mantle, back, lower back and upper rump. Once again, these feather areas were completely green on the birds (as they also were on the two pairs of parent birds), so it had to be the subspecies Tanygnathus lucionensis talautensis. These birds came from an aviculturist who generally maintains a very high level of hygiene and besides has a bird collection that has been clinically tested for the common parrot diseases. Even so, I always also have newly purchased birds re-tested for diseases in connection with the 6-week quarantine period during which they are physically isolated. All the various tests turned out to be negative, but already a few days into the quarantine period the male still seemed to be a bit lethargic and he sneezed once in a while. So, once again he had to be examined by a veterinarian specialized in bird diseases, who took renewed samples from the bird. After laboratory analysis it turned out that the bird had a special rod bacterium that caused sinusitis, so it was put on a penicillin cure with a targeted antibiotic, but it didn't help, the bird got worse and had to remain in quarantine. Another two cures of other and more extensive penicillin preparations over several weeks (with an intermediate period of rebuilding the intestinal flora in the stomach) took place, but nothing helped, it was a chronic sinusitis, and as the bird's condition now was really bad, it also had to be euthanized without at any point having left the quarantine station for several months. The female bird, which is a big beautiful bird in top condition, had no symptoms at any time (sinusitis is not contagious), so this time I decided to keep the surviving female bird.
In 2022 I had the opportunity to buy several young birds from different breeding pairs from the same breeder, who gave me a price reduction on one of the new birds, because I had lost the male that I had bought from him the year before. All these birds came well throughout the quarantine period and besides the fact that they all seem very vital, they also have a very natural behavior, and I have to say that they really make use of their large beaks to gnaw on the many fresh (and unsprayed/uncontaminated) natural branches with which they continuously are supplied. The birds are all, without exception, completely green in the plumage on the mantle, back, lower back and upper rump, and therefore - once again - it just had to be the subspecies Tanygnathus lucionensis talautensis, cf. the information I had received from the various breeders I had spoken to around Europe. Besides, the birds also visually corresponded to a very lifelike drawing in Joseph M. Forshaw's magnificent book, “Parrots of the World” (1st edition, 1973), on page 189, where there is an absolutely excellent color drawing of Tanygnathus lucionensis talautensis made by the outstanding illustrator, the late William T. Cooper (the drawing shows an adult male only with a blue nape (rest of the head is green) and with completely green mantle, back, lower back and upper rump).
The doubt about which subspecies arises
From 2018, I - as several other European aviculturists probably also have heard - were told the story that the subspecies we today have in human care in Europe mostly are the Tanygnathus lucionensis talautensis subspecies, but over time I became more and more uncertain whether this was true.
After acquiring my birds, I later on bought the poster, “Großschnabelpapageien”, from Arndt-Verlag (Thomas Arndt) containing “all” the Tanygnathus species and subspecies, where a pair of Tanygnathus lucionensis talautensis (see photo below) is illustrated with almost completely blue heads in contrast to the adult specimen of this species shown with only a clearly defined blue nape in the color drawing in "Parrots of the World" mentioned above (though Joseph M Forshaw mentions that “blue on crown and occiput variable”, which implies the possibility of a certain form of variance).
Tanygnathus lucionensis talautensis: This photo shows an extract from Arndt-Verlag's poster, “Großschnabelpapageien”, where a male and female of this subspecies are illustrated with almost completely blue heads in contrast to the adult specimen of this species shown with only a clearly defined blue nape in the color drawing in Joseph M. Forshaw’s book, "Parrots of the World".
I was thereafter wondering if it was possible to find a trustworthy and competent person, who actually had been on the Talaud islands and seen how Tanygnathus lucionensis talautensis looks like in wildlife, then it once and for all could be discovered whether the birds that I and other European aviculturists have bought in reality are of this species, or maybe a completely different one.
I then remembered that I during the COVID19-crisis was invited to attend in a Zoom meeting on Wednesday, 22nd September 2021 organized by The Avicultural Society of Australia. Here Mr. Mehd Halaouate held a presentation on the topic "Challenges and Rewards of a mixed Collection". It was an extremely interesting presentation, which included some absolutely stunning photos of wildlife birds from Indonesia including Papua. In many cases the presentation also contained photos of parrot species that we - the European aviculturists - only can dream of and read about in books. During this presentation Mehd Halaouate also showed some photos of birds from the Talaud Islands where he had followed the natural bird life.
In November 2021 I therefore contacted Mehd Halaouate, and asked him, if he during his visits to the Talaud Islands had observed the Tanygnathus lucionensis subspecies and maybe even had taken any photos of this subspecies during his stay. Mehd Halaouate got back to me and stated that he actually had observed Tanygnathus lucionensis talautensis in its natural habitat and also had taken photos of the birds even though he didn't get many since it was challenging to find them and they weren't kept as pets on these islands. I was completely overwhelmed to see Mehd Halaouate’s photos of these birds from the wild and below you can see a couple of these photos of which one actually shows that a big part of the upper head of this subspecies is blue, not just the crown (nape or neck patch). According to Mehd Halaouate most of the birds in the Talaud Islands have heads where almost all the upper parts are clearly blue, and those birds whose heads not almost were entirely blue appeared to be uncolored fledging juveniles as he were there during the breeding season, August and September.
Tanygnathus lucionensis talautensis: This is a photo of an adult, fully colored bird taken by Mehd Halaouate during one of his two trips to Krakelong which is one of the islands that make up the Talaud Islands. Mehd Halaouate was there doing some conservation work for the Red-and-blue lory (Eos histrio).
Tanygnathus lucionensis talautensis: This is also a photo taken on Krakelong Island that shows that the intensity of the blue coloration on the head was not present in all birds. According to Mehd Halaouate it was probably because he was there during August and September which is the breeding season where he encountered many fledging juvenile birds.
Renewed contact with Mehd Halaouate
Time passed and I couldn't help thinking that now that my Tanygnathus lucionensis cannot be of the subspecies talautensis, what kind of subspecies is it then?
In the intervening period I had of course done my homework and made my own research and had come to the conclusion that my birds could probably be Tanygnathus lucionenis salvadorii, which, like Tanygnathus lucionensis talautensis, is the only other (described) subspecies - approved by Howard and Moore - which has a complete green mantle, back, lower back and upper rump.
This subspecies also appears in Arndt-Verlag's poster, “Großschnabelpapageien”, and is shown in the photo below.
Tanygnathus lucionensis salvadorii: This photo shows an extract from Arndt-Verlag's poster, “Großschnabelpapageien”. Apart from Tanygnathus lucionensis talautensis, this is the only other subspecies - recognized by Howard and Moore - which has a complete green mantle, back, lower back and upper rump.
So, in order to get a second opinion and be quite sure about the subspecies determination I decided to contact Mehd Halaouate again to ask him what subspecies he would judge my Tanygnathus lucionensis to be. I sent him various available color descriptions of the relevant subspecies as well as - not least - photos of my birds, all of which can be seen excerpted below.
Descriptions of the Tanygnathus lucionensis subspecies
As you know, the world's leading current nomenclature/taxonomy, “THE HOWARD AND MOORE COMPLETE CHECKLIST OF THE BIRDS OF THE WORLD” mentions up to 4 subspecies (hybridus, talautensis, salvadorii and horrisonus) in addition to the nominate subspecies, but unfortunately this work does not contain any descriptions of the individual subspecies. Instead, one has to look in different zoological museums around the world to find stuffed birds of the so-called "holotype specimens" for each subspecies, and this is quite an unmanageable task. I therefore had to find other trustworthy sources that contain color descriptions of the various subspecies:
Source 1 for color description: Joseph M. Forshaw
As already mentioned, it is very unfortunate that there is no credible, scientifically evidence-based literature covering the entire Tanygnathus genus, with correct and detailed descriptions of each subspecies. In the absence of this, I instead rely on the Australian ornithologist Joseph M. Forshaw's impressive book “Parrots of the World”, which however, only mentions 2 subspecies (hybridus and talautensis) besides the nominate subspecies, but the description of the two subspecies here is unfortunately not adequate. However, in the original edition of this book (1st edition, 1973), on page 189, there is an absolutely excellent color drawing of Tanygnathus lucionensis talautensis made by the outstanding illustrator, the late William T. Cooper (an adult male only with a blue nape (not on the rest of the head) and completely green mantle, back, lower back and rump).
Source 2 for color description: Thomas Arndt (Arndt-Verlag)
In the well-known "Lexicon of Parrots" (version 3.0), Thomas Arndt mentions the same two subspecies as mentioned by Joseph M. Forshaw (hybridus and talautensis).
However, in his poster, “Großschnabelpapageien”, containing species and subspecies from the Tanygnathus genus, Thomas Arndt shows both the nominate subspecies of Tanygnathus lucionensis and 3 subspecies (hybridus, talautensis and a “new” one, namely salvadorii). If one looks closer at this poster from Thomas Arndt (see above), he has illustrated Tanygnathus lucionensis talautensis - both the male and the female - with almost completely blue heads in contrast to the color drawing in "Parrots of the World" (also mentioned above) that shows an adult Tanygnatus lucionensis talautensis with only a blue nape patch and not a blue head. This corresponds to the photos Mehd Halaouate himself took of adult birds of this subspecies during his two visits to the Talaud Islands.
However, and very interesting, on the same poster from Thomas Arndt, he also displays the subspecies Tanygnatus lucionensis salvadorii that in particular deserves attention. This subspecies is shown with only a blue nape and completely green back and rump (and looks very much like the Tanygnatus lucionensis talautensis, that is shown in Joseph M. Forshaw’s book mentioned above).
Source 3 for color description: World Parrot Trust
What is very remarkable is that the World Parrot Trust, on its website - like Joseph M. Forshaw - only mentions the subspecies hybridus and talautensis, but when one read further in the text under the actual description of the two subspecies, World Parrot Trust actually also describes salvadorii under the description of talautensis (they mention that the two different subspecies look the same). How this is possible, I can't quite see through.
Summary overview of visual differences between the individual subspecies
Since there is no adequate and sufficient information on the sizes (length and weight) of the individual subspecies, only selected visual special characteristics for the plumage (color differences) of the individual Tanygnatus lucionensis subspecies are given below:
Note to the above table:
How the fifth subspecies - Tanygnathus lucionensis horrisonus, mentioned by Howard and Moore - looks like, I do not know in details. However, if you read “PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY BULLETIN 13, “Notes on a Collection of Birds from Mindoro Island, Philippines” by S. Dillon Ripley (Peabody Museum of Natural History, Yale University) and D. S. Rabor (Biology Department, Silliman University), which was published 31st December 1958 (New Haven, Connecticut), you can read on page 36, that the authors after carefully studies consider “horrisonus” as synonyms of “lucionensis”, which means that it must have blue mantle, back, lower back and upper rump, and my birds don't have these hallmarks. According to the newest version of Howard and Moore “horrisonus” is still recognized as an independent subspecies.
I therefore assume that my birds must be either Tanygnathus lucionensis talautensis or Tanygnathus lucionensis salvadorii, as both of these subspecies have green backs and green upper rumps.
Personally, I strongly believe that my birds are Tanygnathus lucionensis salvadorii, as I have seen the various adult parents of the birds that I have bought and none of them have a nearly completely upper blue head, but only a blue patch on the nape.
Furthermore, my birds don’t have any blue coloration - nor bluish tinge - on the mantle, back, lower back and the upper rump, and to my knowledge, both the nominate subspecies, Tanygnathus lucionensis lucionensis, and the subspecies, Tanygnathus lucionensis hybridus, clearly have these hallmarks.
I rule out that my birds are crossed with other subspecies, as they have no blue at all on the back or rump, nor do they have more than a clearly defined blue nape patch.
Some of the photo documentations sent to Mehd Halaouate
I sent several photos of my birds to Mehd Halaouate and some them are showed below. All my birds are from last year (2022), except from the bird in front on photo 09 and 13, this bird is from 2021.
The photos of my birds are unfortunately not of the best quality, as they were taken with the zoom-function on my mobile phone and have become somewhat grainy. I should of course also have captured some of the birds and taken photos of them from the front and from the back with spread wings, but since these birds live in aviaries close to some other parrot species that already have laid and are incubating eggs, it was unfortunately not possible to take some better photos for the near future. Here are some photos of my birds:
After having read the above descriptions and studied the forwarded photos of my birds, Mehd Halaouate commented that my birds looked stunning and very healthy and furthermore mentioned that in his opinion they look more like Tanygnathus lucionensis salvadorii. At the same time, one must of course bear in mind that there are color variations among birds in the wild too that can make establishing the right distribution of the bird pretty impossible.
However, according to Mehd Halaouate most of the subspecies that were kept during his time as breeder in Europe, they were coming from Mr. Antonio de Dios in the Philippines (owner of “Birds International Inc.”, a large private parrot breeding farm in the Philippines), and they mostly were Tanygnathus lucionensis salvadorii. The for many aviculturists well-known Antonio de Dios exported a huge number of unique species to Europe especially to Holland and Belgium. Some of the other species he sent were species like Fig parrots, a big number of lories and lorikeet species, Great-billed parrots (Tanygnatus megalorynchos) and Moluccan king parrot subspecies.
Mehd Halaouate also mentioned that he had to search for some of his earlier photos of the Tanygnathus lucionensis talautensis to definitively prove how the adult birds of this subspecies looks like in wildlife, some truly unique photos, a few of which can be seen below. It is photos of a pet bird taken at a bird trader in Sulawesi (since the local people in Talaud Islands don’t keep this bird as a pet) and it does INDEED have intensive blue head covering all over the upperparts of the head and even around the eyes.
Tanygnathus lucionensis talautensis: I have never seen anything like it. This amazing close-up photo shows an adult and fully colored Tanygnathus lucionensis talautensis. The photo clearly shows how most adult birds of this subspecies that live in the wild on the Talaud islands look like i.e., where the blue plumage does not just form a clearly defined neck spot, but instead is spread over most of the upper part of the whole the head. Young birds do not have the same distribution of the blue color in the head. It is a pet bird that sits with a bird dealer on the island of Sulawesi. Photo: Mehd Halaouate.
Tanygnathus lucionensis talautensis: This equally amazing close-up photo shows the above pet bird of Tanygnathus lucionensis talautensis, just seen from a different angle. For some reason, the local inhabitants of the Talaud Islands do not keep this species as a pet bird, so this individual is photographed on the island of Sulawesi, which is located southwest of the Talaud Islands. The Talaud islands actually make up the area that is called North Sulawesi. Photo: Mehd Halaouate.
According to Mehd Halaouate the poster, “Großschnabelpapageien”, from Arndt-Verlag with the color drawing of Tanygnathus lucionensis talautensis is a bit exaggerated with the blue coloration of the head, throat and neck. Mehd Halaouate have never seen any bird with that much blue on the head.
Unexpected challenges in rewilding Tanygnathus parrots
During my dialogue with Mehd Halaouate, he also mentioned the following - for me highly surprising - problem, which is linked to the rewilding of the Tanygnathus parrots, which has been confiscated as a result of smuggling:
A factor which will make things even worse for establishing “the right ID” of many parrots in the wild in the near future is the fact that the forestry department in Indonesia does not have sufficient knowledge about differentiating between subspecies and in some cases even species. They tend to release confiscated parrots from the illegal trade in the wrong distribution area (habitat). As a former manager in World Parrot Trust Mehd Halaouate has had to stop a few of these releases because the parrots did not belong in specific islands they were meant to be released at. The authorities, unfortunately, had managed to do the damage thus these actions will cause the inter- or cross-breeding between the subspecies which will result in new colour variations. The worst result of this is that some subspecies in the nature will lose their purity.
This is an angle that I have not thought of before, and the authorities in the affected countries simply have to try to find a sustainable solution to the problem, so that before rewilding takes place, quality assurance must always be carried out to ensure that the birds are belonging to the birds' original distribution area (habitat), which presupposes a further educational and competence-related upgrade of the authorities' employees.
Also from this perspective, it is very important that we - the serious aviculturists (breeders) - are trying as much as possible to keep our birds pure as these hopefully one day will assist the wild populations. According to Mehd Halaouate the pace in which the illegal trapping and the wildlife trade is proceeding right now, many parrot species, for their survival, will rely on the captive populations, so we - the aviculturists (breeders) - must act responsible, professional and only work with pure birds at the subspecies levels.
I have on earlier occasions tried to initiate a dialogue about the issue of subspecies determination of this genus with some other European aviculturists who keep these birds, but up until now it had seemed that no one really was interested in getting to the bottom of this issue, so they generally just call their birds for Blue-naped Parrots (Tanygnatus lucionensis). I always try to keep my different parrots pure at the subspecies level (except from monotypic taxon) and I always only breed specimens of the same subspecies with each other, and I would strongly encourage other responsible breeders to do the same.
This was the story of not always believing everything you hear from other aviculturists/breeders. It is good to have a healthy skepticism and to try to form your own opinion about things. So, the next time I hear an aviculturist/breeder saying that the subspecies we have the most of in Europe is Tanygnathus lucionensis talautensis, I can now for sure say that it is not true and instead tell them a completely different story.
I hope that interested readers have found pleasure in reading this article about an exciting parrot species that there is rarely written many lines about in parrot books and in avicultural magazines.
A great thank you to Mehd Halaouate for having contributed to this article with several unique photos and detailed information about the subspecies of Tanygnathus lucionensis.
Tanygnathus lucionensis talautensis: I could not resist ending up with showing this close-up profile photo of the magnificent pet bird of this subspecies. I just keep on turning back to study the photos of this very special subspecies with a nearly completely blue head. Photo: Mehd Halaouate.