Falcons & Parrots

The 46 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

PASSERIMORPHAE

There are several lines of evidence supporting this clade. First are the conventional DNA analyses of Ericson et al. (2006) and Hackett et al. (2008). Although these use the controversial 7th intron of the β-fibrogen gene, the clade also appears in Ericson et al.'s supplementary ESM-6 tree, which does not use the gene. Wang et al. (2012) use 30 nuclear introns (not including any introns of β-fibrogen). Their work also supports the notion that passerines and parrots are sisters, and that the falcons and seriemas are their closet relatives. Nabholz et al. (2011) also support the close relation between the passerines and parrots, although taxon sampling is very sparse and they do not consider the falcons. Passerimorphae also appear as a clade (under the name Eufalconimorphae) in Suh et al. (2011). They use the completely different approach of retroposon insertions to investigate avian phylogeny. They also note the similarities in the vocal apparatus and the fact that both parrots and many passeriformes learn their song. Mayr (2011) has also noted that the Passeriformes have an extinct sister group, Zygodactylidae, that indicate a common heritage with the zygodactyl parrots.

CARIAMIFORMES Wagler, 1830

Cariamidae: Seriemas Bonaparte, 1850 (1836)

2 genera, 2 species HBW-3

The seriemas have sometimes been considered Gruiformes, but genetic evidence points elsewhere. Hackett et al.'s (2008) analysis found them sister to the remaining three orders, while Ericson et al. (2006a) suggest they belong in the Falconiformes. Either way, they are next in the list!

Although there are currently only two species of Seriemas, they are a remnant of a much larger group, the Cariamae. Like the Cariamidae, these have traditionally been placed in the Gruiformes. Besides the Cariamidae, the Cariamae are thought to have included three now-extinct families: the Phorusrhacidae, Bathornithidae and Idiornithidae. The Phorusrhacidae are the well-known “terror birds” of ancient South America. See Alvarenga and Höfling (2003) for more on the Phorusrhacidae.

FALCONIFORMES Sharpe, 1874

Falconidae: Falcons, Caracaras Leach, 1820

12 genera, 67 species HBW-2

Click for Falconidae tree
Click for Falconidae tree

The phylogeny is now based on the multigene analysis of Fuchs et al. (2014), who considered almost all of the species. Fuchs et al. (2014) calibrate their tree based on two fossils. They found that crown-group Falconidae date to the Oligocene. This is interesting as the crown-groups of their sister taxa (Psittaciformes plus Passeriformes) are older, much older. If the falcons are correctly placed as sister to the parrots and passeriformes, they must have originated in the early part of the Eocene or more likely Paleocene, perhaps 50-60 million years ago (some estimates make them even older). Yet only one lineage from the Oligocene (roughly 30 million years) is the ancestor of all the extant Falconidae, with all other Falconidae lineages from the previous 20-30 million years becoming extinct.

Previously, I'd used the papers by Griffiths (1999) and Griffiths et al. (2004) for the basic taxonomy of the Falconidae. This had been supplemented by Fuchs et al. (2011b) for the forest-falcon clade (Herpetotherinae), and Fuchs et al. (2012a) for the caracaras (Caracarinae). Many of the caracaras are closely related, and it would not be unreasonable to merge Ibycter, Milvago, and Phalcoboenus into Daptrius.

Fuchs et al. (2014) found that the two species of Polihierax are not closely related, with the Pygmy Falcon closer to Microhierax and the White-rumped Falcon nearer Falco. Accordingly, the White-rumped Falcon is placed in the monotypic genus Neohierax (Swann, 1922). Falco itself seems to have originated much more recently, with a common ancestor most likely between 5 and 8 million years ago, a good reason to leave all of Falco in a single genus. In contrast, the Neohierax/Falco split was likely 10-15 million years ago. To further emphasize the division in Falconinae, I've recognized two tribes within it: Polihieracini and Falconini.

The African kestrel clade was studied by Groombridge et al. (2002). Based on their work, Rock Kestrel must be split from Eurasian Kestrel because the Australian Kestrel is split. The Hierofalco complex was examined in detail by Wink et al. (2004a). In spite of sampling many individuals, they were unable to produce a clean phylogeny. Although this may mean some of the these species should be lumped, they believe the confused situation is due to ancient hybridization. Further analysis is required to sort out the Hierofalco complex. This remains true even after Fuchs et al.'s (2014) analysis. The order of Falco itself is now based on Fuchs et al. (2014) rather than Wink and Sauer-Gürth (2004).

Herpetotherinae: Forest-Falcons Lesson, 1843

Caracarinae: Caracaras d'Orbigny, 1837

Falconinae: Falcons Leach, 1820

Polihieracini Peters, 1931

Falconini Leach, 1820

PSITTACIFORMES Wagler, 1830

Ericson et al. (2006a) suggest the parrots are sister to the passeriformes, as do Hackett et al. (2008).

The overall structure of the Psittaciformes follows Schirtzinger et al. (2012), Schweitzer et al. (2010), and Wright et al. (2008). Christidis and Boles (2008) contains detailed information on Australian parrots. Brown and Toft (1999) and White et al. (2011) provided further information concerning the Cockatoos; Groombridge et al. (2004) and Kundu et al. (2012) were helpful with the Psittaculini; the placement of the Ground and Night Parrots is inspired by Leeton et al. (1994). Tavares et al. (2006) gives a nice overview of the Arini, with Kirchman et al. (2012) providing additional detail. The papers by Ribas et al. (2005, 2007a, b) were helpful concerning the basal Androglossini, while Russello and Amato (2004) was consulted concerning the Amazona parrots. Russello and Amato found that the for last four (possibly five) Amazona species, the genetic data does not seem to match existing subspecies.

The whole order is summed up in the following tree diagram.

Psittaciformes
Click for genus-level tree for Psittaciformes

Strigopidae: New Zealand Parrots Bonaparte, 1849

2 genera, 4 species Not HBW Family

I had earlier used the name Nestoridae, which seemed to be the most common usage for a family containing both Strigops and Nestor. However, both family-group names were introduced at the same time (Bonaparte, 1849), and they have normally been kept separated enough that the issue of priority doesn't arise. I don't have access to Bonaparte (1849), but Bock (1994) does not mention priority between them, listing them as Strigopinae and Nestorinae. Christidis and Boles (2008) also state they have equal priority. Assuming that is correct, the determination of priority falls to the first reviser. That's Bonaparte himself the following year, when he gave priority to Strigopidae by listing Nestorinae as a subfamily (1850, Vol. 1, p. 8).

Cacatuidae: Cockatoos G.R. Gray, 1840 (1825)

7 genera, 21 species HBW-4

The order within the cockatoos is based on White et al. (2011), with some help from Brown and Toft (1999). White et al. (2011) found that the Cockatiel was basal, and I follow their mulitgene analysis (as well as traditional thinking). Note however that Brown and Toft found a different position for the Cockatiel, which also received limited support from Wright et al. (2008).

Three of the black-cockatoos have been moved to the genus Zanda (Mathews 1913, type bauinii) due to substantial separation between them and the Calyptorhynchus black-cockatoos. White et al. (2011) estimate the most recent common ancestor at about 15 million years ago. Interesting, the current Zanda diversity appeared very recently, whereas the split between the two Calyptorhynchus is of much longer standing.

Psittacidae: African and American Parrots Rafinesque, 1815

37 genera, 171 species HBW-4

Schirtzinger et al. (2012), Schweizer et al. (2010) and Wright et al. (2008) identify 5 deep clades in the the remaining parrots: the vasa-parrots, Pesquet's Parrot, an Old World/Australasian clade (Psittaculinae), an African clade (Psittacinae), and an American clade (Arinae). Pesquet's Parrot and the vasa parrots seem to be each other's closet relatives, and we unite them in the subfamily Psittrichasinae. The appear to be sister groups, as do the Psittrichasinae and Psittaculinae, leaving us with two major clades: Psittacidae (Psittacinae and Arinae) and Psittaculidae (Psittrichasinae and Psittaculinae).

How deep are the five clades? Wright et al. (2008) gave two different timelines. In one, they date from the late Cretaceous to the early Paleocene, roughly 65 million years ago. They also provide a later dating, in the Eocene, about 40 million years ago. The earlier dating suppose that the New Zealand Parrots date from the separation of New Zealand from Gondwana, over 80 million years ago. The later dating puts the split of the New Zealand Parrots about 50 million years ago, in the Eocene. Joseph et al. (2012), who seem to prefer an early dating, recommend ranking Psittacinae, Psittrichasinae, and Psittaculinae as families rather than subfamiles. Personally, I prefer the later dating as it fits better with the fossil record. I also follow Howard and Moore 4th ed. and SACC by using only two families: Psittacidae and Psittaculidae.

Of the two main clades, Psittaculidae is slightly larger, so we put it last in the linear order.

Psittacinae: Afrotropical Parrots Rafinesque, 1815

Arinae: Neotropical Parrots G.R. Gray, 1840 (1825)

The Neotropical parrots, Arinae, fall into 5 subclades: Amoropsittacini, Brotogerini, the parrot clade Androglossini, Forpus parrotlets (Forpini), and a narrowly circumscribed conure/macaw clade Arini. The relationships between them are not fully understood at this time—the various relevant papers give conflicting topologies (de Kloet and de Kloet, 2005; Tavares et al., 2006; Wright et al., 2008, Schirtzinger et al., 2012). However, the Tavares et al. (2006) topology seems to fit best with the other papers, and I have adopted it.

In the Tavares et al. topology, Amoropsittacini is the basal group. Schirtzinger et al. (2012) also included Psilopsiagon in their analysis, putting it here in the Amoropsittacini. The Brotogerini appear to be affiliated with the Androglossini, and the Forpini with the core Arini.

Amoropsittacini Brereton, 1963

Brotogerini Wolters, 1975

The ordering of the Brotegerini is based on Ribas et al. (2009). They found that the Brotogeris parakeets fall into two groups of 4 species each. Not surprisingly, the genetic distance between chiriri and versicolurus was fairly small, with the amount being consistent with treatment either as separate species or subspecies of a single species.

Androglossini Sundevall, 1872

The series of papers by Ribas et al. (2005, 2007a, b) and Russello and Amato (2004) have done much to clarify the situation in the Androglossini. The parrots from Pionopsitta to Pyrilia were studied by Ribas et al. (2005). The ordering of the Pionus parrots reflects the results of Ribas et al. (2007a). The results also support a possible split of White-capped Parrot, Pionus seniloides. The Hapalopsittaca order is based on Quintero et al. (2013).

Yellow-headed Complex The big issue here is the Yellow-headed Amazon complex—the last six species in the list. This group has long been controversial, and although the taxonomy here is close to that of AOU, it seems to be incorrect. The papers by Rusello et al. (2004) and Eberhard and Bermingham (2004) showed the Turquoise-fronted Amazon is a member of this complex. Ribas et al. (2007b) sampled many more of the South American birds in an effort to clear up the mysteries of this group, but the relationships there remain confusing. Urantówka et al. (2014) helped clarify matters a bit by including A. barbadensis.

As things currently stand, the species of the Yellow-headed complex are divided into 15 subspecies: A. aestiva aestiva, A. aestiva xanthopteryx; A. auropalliata auropalliata, A. auropalliata caribaea, A. auropalliata parvipes; A. barbadensis, A. ochrocephala nattereri, A. ochrocephala ochrocephala, A. ochrocephala panamensis, A. ochrocephala xantholaema; A. oratrix “guatemalensis” (not formally described), A. oratrix belizensis, A. oratrix hondurensis, A. oratrix oratrix, and A. oratrix tresmariae. However, DNA tells a different story. I'm not entirely sure how the subspecies fit together as the parvipes clade and tresmariae were not included in the same paper, so I leave their relative position unresolved.

Most of the A. ochrocephala subspecies are very closely related to A. aestiva, including the A. o. ochrocephala from the lower Amazon (“ochrocephala”). Based on current information, these are too close and too mixed up to meaningfully separate. It makes sense to put them all into A. aestiva. However, the ochrocephala from Colombia and Venezuela are a different story. These birds appear to be basal in the Yellow-headed/Turquoise-fronted group. I think the ochrocephala type specimen is from Venezuela, so these birds keep their name (no nasty quotes around it). The Middle American subspecies currently in oratrix and auropalliata are fairly closely related, and could be reasonably grouped under A. auropalliata (auropalliata Lesson, 1842 has priority over oratrix Ridgway, 1887). That gives us four species as shown in version one (the species names are in red). Some have argued that the Tres Marias Amazon should be considered a separate species, and version II shows how such a tree would look under the assumption that parvipes is closer to oratrix than to tremariae. The other Middle American subspecies seem to form monophyletic clades. If interbreeding is sufficiently limited, they too could be promoted to species status. Yet another option is to put them all into one species which would take the name A. aestiva, not A. ochrocephala.

The list currently follows IOC in recognizing the Tres Marias Amazon, Amazona tresmariae, formerly a subspecies of Amazona oratrix. This also implies that the Panama Amazon, Amazona panamensis should be separated too, and I have done so. The situation with auropalliata and oratrix seems to leave auropalliata paraphyletic. I prefer to wait for more information before resolving this issue.

Wenner et al. (2012) found a substantial amount of genetic structure in the Mealy Amazon, Amazona farinosa. The Central American populations to west Panama (guatemalae and virenticeps) formed a separate clade from those in the Darien and South America (inornata and farinosa/chapmani), with 3.5-5.4% genetic distance in cytochrome-b. Such differences often indicate separate species. However, they did not look in detail at the situation within Panama where the two clades are in contact. Until such a study is done, I will treat them as part of a single species as there doesn't seem to be any separating mechanism, which suggests that only one biological species is involved. Contrary to its treatment in recent checklists, the Choco-Darien form inornata was clearly distinct from farinosa/chapmani and should at least be recognized as a subspecies.

Forpini: Forpus Parrotlets Brereton, 1963

The arrangement within Forpus is based on B.T. Smith et al. (2013). The species limits may need some adjustment. In particular, the Blue-winged Parrotlet, Forpus xanthopterygius, may not be monophyletic.

Arini: Conures and Macaws G.R. Gray, 1840 (1825)

Tavares et al. (2006) found strong support for a basal Deroptyus-Pionites grouping, sister to the remaining Arini. This arrangement also got mixed support from Wright et al. (2008), and is adopted here.

The Yellow-eared Parrot, Ognorhynchus icterotis, doesn't seem to have been included in any genetic studies. It probably groups with the remaining Arini, but it's true position is unresolved.

Rhynchopsitta may group with Pyrrhura (Tavares et al., 2006), or maybe not (Kirchman et al., 2012). The arrangement of the Pyrrhura parakeets is based on Ribas et al. (2006).

The southern genera Enicognathus, Cyanoliseus, and Anodorhynchus seem to branch successively, bringing us to the macaws, Aratinga parakeets, and allies. This group includes a well-supported macaw clade, Cyanopsitta though Ara, and a small, well-supported grouping of Leptosittaca, Diopsittaca, and Guaruba. Urantowka et al. (2013) found that the Blue-crowned Parakeet is sister to Diopsittaca. I've placed it in the monotypic genus Thectocercus (Ridgway 1912).

The traditional version of Aratinga is not a monophyletic group (Ribas and Miyaki, 2004). The problem is how to separate them, and how they fit with the other two groups. Recently, Kirchman et al. (2012) included enough species in their analysis to split up Aratinga in a reasonable way. Support for the overall phylogeny, as followed here, is mediocre. I think the division of Aratinga is on more solid ground.

Kirchman et al. show three clades containing Aratinga parakeets. One clade is sister to the Carolina Parakeet, Conuropsis carolinensis. It includes the type species of Aratinga, the Sun Parakeet, Aratinga solstitialis and also auricapillus. Ribas and Miyaki (2004) showed that jandaya is sister to auricapillus, so it belongs here too, as does the the recently split maculata (see Silveira et al., 2005. Note that the scientific name has been corrected to A. maculata from A. pintoi by Nemésio and Rasmussen, 2009). Kirchman et al. show the Nanday Parakeet (usually Nandayus nenday) as the basal member of the group. However, a look at Ribas and Miyaki suggests that the Dusky-headed Parakeet, Aratinga weddellii, is the true basal member. Accordingly, I've merged Nandayus into Aratinga, which applies to these dark-billed parakeets. The whole lot may be sister to the macaw clade.

The horn-billed, green-winged, generally red and green “Aratinga” also appear to form a clade. The oldest available genus name is Psittacara (Vigors 1825, type leucophthalmus). Kirchman et al. found the Psittacara sister to the Leptosittaca-Guaruba group.

The other clade of former Aratinga includes aurea, nana, and pertinax. Comparison with Ribas and Miyaki suggests that cactorum belongs here too. Since canicularis is thought to form a superspecies with aurea, it also goes in this group. The oldest available name seems to be Eupsittula (Bonaparte 1853, type canicularis). Kirchman et al. show this clade as sister to Rhynchopsitta. Maybe, but stronger evidence from other papers (e.g., Tavares et al., 2006) put Rhynchopsitta elsewhere, and so do I.

Psittaculidae: Old World Parrots Vigors, 1825

46 genera, 191 species HBW-4

Psittaculidae contains two subfamilies: Psittrichasinae and Psittaculinae.

Psittrichasinae: Pesquet's & Vasa Parrots von Boetticher, 1959 (1854)

Kundu et al. (2012) examined a variety of Indian Ocean parrots. They found that that extinct Mascarene Parrot, Mascarinus mascarin, belongs in the same genus as the vasa parrots. As Mascarinus (Lesson, 1830) is an older name that Coracopsis (Wagler, 1832), the entire group takes the genus name Mascarinus. The subfamily name remains unchanged. Kundu et al. also found that the Seychelles Black Parrot, usually considered a race of the Lesser Vasa Parrot, is fairly distant from the Lesser Vasa Parrot. Accordingly, I treat it as a separate species, Mascarinus barklyi.

Psittaculinae Vigors, 1825

Schweizer et al. (2010) and Wright et al. (2008) both found 5 major clades within the Psittaculinae, although neither group included Pezoporus or Psittacella in their analysis.

So where does Pezoporus go? Based on Leeton et al. (1994) and Miyaki et al. (1998), it seems to be sister to Neopsephotus + Neophema, and I treat all three as a single clade (Pezoporini). However, this is not as solid as it might be because neither included the Agapornithini in their analysis, which Wright et al. had as sister group to Neopsephotus + Neophema. More recently, Joseph et al. (2011) included all the relevant taxa in a multigene analysis and established that Pezoporus is sister to Neopsephotus + Neophema.

Joseph et al. (2011) also showed that the Psittacella tiger-parrots form a separate clade. This gives us 6 clades: Psittaculini; Psittacellini; Platycercini; Pezoporini; Agapornithini; Loriini.

The Psittaculini are basal to the others. This tribe contains three main pieces, the pygmy parrots (Micropsitta), Alisterus through Polytelis, and Prioniturus through Psittacula, possibly excepting the extinct Lophopsittacus and Necropsittacus. These two, which might be each other's closest relatives have long been believed to be related to Psittacula. However, it is possible that they are closer to the Mascarene Parrot and Vasa Parrots. There is some question about the monophyly of Polytelis, although Joseph et al. (2011) and Schweizer et al. (2010) have different takes on the nature of the problem. The treatment of the Prioniturus racket-tails is based on Schweitzer et al. (2012). They found that the race mindorensis is not part of P. discurus, but is more closely related to P. platenae. Both plumage and genetic distance suggest it is a separate species. I could find an existing English name, so I'm referring to it as Mindoro Racket-tail.

Joseph et al. (2011) show that the Psittacellini tiger-parrots are sister to the remaining 4 tribes. Exactly how those 4 tribes fit together remains somewhat uncertain. I'm following the Schweizer et al. (2010) topology, which is consistent with the MP analysis of Wright et al. (2008), and figure 2 of Joseph et al. (2011). Joseph et al. argue that this result, from a restricted data set, is more likely to be correct. Their figure 1 and the Bayesian analysis in Wright et al. give a different arrangement.

Platycercini has two parts: Psephotus through Platycercus and Lathamus through Cyanoramphus. All the genera of the first group are sometimes lumped in an expanded Platycercus. Only the uncertainty about whether the Pezoporini are sister to this group, prevents me from including them in the Platycercini as in Collar (1997). It appears that the traditional Psephotus is not monophyletic (see Joseph et al., 2011; Schweizer et al., 2010, 2013), with Red-rumped Parrot, Psephotus haematonotus, separated from the rest. As it is the type species for Psephotus, we go to the next available name for the other birds formerly placed in Psephotus. That name appears to be Psephotellus (Matthews 1913, type pulcherrimus). I'm following a blend of Schweizer et al. (2013) and Joseph et al. (2011) for this branch of the Platycercini. Although they are clearly part of the clade, the placement of Northiella and Psephotus remains uncertain. They show up in slightly different locations in different analyses.

Next we consider the Loriini. Cyclopsitta/Psittaculirostris is the basal group in the Loriini. The other clade starts with Melopsittacus.

Now that Bird Life International has joined IOC in splitting the Rainbow Lorikeet, Trichoglossus haematodus, into three species, I'm adopting that split. BLI based their split is based on the Tobias et al. (2010) criteria for allopatric species, which is a reasonable rule of thumb in the absence of conflicting evidence. However, as BLI has not yet released a detailed analysis, I will not generally adopt these splits. In this case, I make an exception because IOC had already adopted them. The three species are:

Psittaculini Vigors, 1825

Psittacellini: Tiger-Parrots Wolters, 1975

Joseph et al. (2011) found that the Tiger-Parrots were not closely related to any of the remaining parrot tribes, but formed a branch sister to all of them.

Pezoporini: Ground and Night Parrots Bonaparte, 1838

The Ground Parrot, Pezoporus wallicus, has been split into Western Ground Parrot, Pezoporus flaviventris, and Eastern Ground Parrot, Pezoporus wallicus. See Murphy et al. (2011).

Platycercini Selby, 1836

The genus name Prosopeia has been replaced by Pyrrhulopsis, which has priority. See Gregory and Dickinson (2012).

Agapornithini Salvin, 1882

Loriini: Lories Selby, 1836

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