Ratites and Tinamous

The 47 Orders














The TiF Checklist

The checklist can be viewed in two ways. You can either view the annotated checklist on these web pages or download a list. The downloadable lists are Excel csv files that can be imported into spreadsheets such as Excel, or easily manipulated by programs such as perl. Four lists are available in csv format. The ABA list includes only ABA recognized species, but in TiF order, with TiF families. The South American list uses TiF species rather than SACC species.

Further, Stephen Nawrocki has provided an excel version of the worldlist, version 2.79.

You can view an annotated version by clicking on the list of bird orders on the right, or by going to the family index, or by clicking on the family names in the various tree view pages. In the annotated list, recently extinct species and species whose taxonomic placement is particularly uncertain are color-coded. In some cases, superfamilies, subfamilies, tribes, and other groups have been added to help show how the birds are related.

The 47 Orders

The TiF checklist currently groups the birds in 47 Orders and 251 families. Both a order-level and family-level trees are now availalbe in pdf format. Due to its length, the family tree is split into 5 parts.

Click for order-level tree Click for family-level tree
47 Orders 251 Families

The 47 orders of the TiF list include a few more orders than most other lists. However, I wanted to make sure that each of the orders was monophyletic, meaning that they include all descendants of a common ancestor. I did not have confidence that a shorter list would do. As it currently stands, I am highly confident that each order is monophyletic.

Supra-ordinal Groups

I am also highly confident in 12 supraordinal groups (including 4 subgroups) that are supported by Jarvis et al. (2014), Prum et al. (2015), and Suh (2016). Except for Otidimorphae, Eurypygimorphae, and Ardeae, these are also supported by Hackett et al. (2008). Figure 1 in Suh (2016) gives more detail on support for these groups. Not all of the papers include all 47 TiF orders. They are listed below, along with 3 orders that do not belong to any of the groups.























Independent Orders


Higher Phylogeny

The basic problem of higher phylogeny has now been reduced to the appropriate placement of the 8 high order groups and 3 independent orders.

The division into Palaeognathae and Neognathe (all other living birds) was first proposed by Pycraft (1900). Sibley, Ahlquist, and Monroe (1988) suggested that the Galloanserae and Palaeognathae were basal lineages among the living birds (Neornithes) and united them as Eoaves, with all other birds grouped as Neoaves. Sibley and Ahlquist (1990) recognized that the Palaeognathae and Galloanserae were separate branches, and restricted Eoaves to be identical to Palaeognathae. Since then, the evidence in favor of this arrangement has only grown stronger. The names Palaeognathae and Neognathae are due to Pycraft (1900), while Neoaves comes from Sibley, Ahlquist, and Monroe (1988). The Galloanserae are included in Neognathae, but excluded from Neoaves.

This leaves us with 6 major supra-ordinal groups and 3 independent orders to classify. Here, we run into a big problem. Jarvis et al. (2014), Prum et al. (2015), and Suh et al. (2015) have given somewhat inconsistent results concerning how these 9 groups relate. Indeed, we have been seeing such inconsistencies for some time. Thus both Brown et al. (2008) and Morgan-Richards et al. (2008) informed us that the Metaves hypothesis was incorrect. However, their own preferred topologies would likely have been rejected by each other's data, and have since fared no better than Metaves.

The Polytomy: The Hoatzin Problem on Steroids

Suh (2016) has argued that these inconsistencies stem from a hard polytomy at the base of Neoaves involving all 9 groups, a polytomy that cannot be satisfactorily resolved with current data and methods of analysis.

In alphabetical order, the nine troublesome taxa are: Ardeae, Charadriiformes, Columbimorphae, Gruiformes, Mirandornithes, Opisthocomiformes, Otidimorphae, Strisores, and Telluraves.

So how should we arrange this hard polytomy? If Suh (2016) is correct, any arrangement will be pretty arbitrary. That's what a hard polytomy means. One way to handle it is to order them from least diverse to most diverse. On the TiF list, this means ordering them by species diversity rather than by diversity of the next lower major group. The reason for this is that the groups between the group we are ordering and species are quite arbitrary. They mean different things in terms of time depth and genetic distance depending on where you are in the tree. Biological species have a greater tie to biological reality than do these other groups, even though that tie is still fairly loose. So I count species in each group at a given level to order them.

If I ordered them by species diversity, the order would be Opisthocomiformes, Mirandornithes, Gruiformes, Otidimorphae, Columbimorphae, Ardeae, Charadriiformes, Strisores, Telluraves.

However, I think we can do a little better. I don't think there's zero information about the polytomy. Indeed, looking at Figure 2 of Suh (2016), it appears that the Columbimorphae (which are probably, but not certainly monophyletic) are closest to the root and the Otidimorphae are probably next. The Strisores may come next. After that, things are murkier. Indeed, these groups often show up near the root of Neoaves in other analyses.

That reduces the problem to 6 taxa: Opisthocomiformes, Mirandornithes, Gruiformes, Ardeae, Charadriiformes, and Telluraves. How should we then order these? I've played around with different arrangements, trying to group waterbirds and landbirds separately, or with other intuititive groupings. In the end, I decided it was better to acknowledge our lack of knowledge.

I've ordered the remaining six taxa by size. In the TiF list, size means number of species. We could use the number of orders, or families, or genera, but I like to use number of species. The reason for this is simple. The number of species is less arbitrary—we have a biological criterion for species status. The other concepts are more arbitrary, and too often reflect the preferences of a particular ornithologist or committee.

What Caused the Polytomy?

The likely cause of the hard polytomy lies with the end-Cretaceous extinction. The scenario is that a single neoavian lineage survived (as did at least one lineage each of Galloanseres and Palaeognathae). The single lineage soon evolved into many lineages to cope with the new, post-apocalyptic ecology. The ancestors of the 9 groups diversified too quickly for it to be reflected in the genes (incomplete lineage sorting) and may have also have continued to hybridize. This makes the genetics of the splits difficult or even impossible to sort out. It is certainly beyond what we can do in 2016.

When thinking about the polytomy, you should keep in mind that the ancestors of the nine groups would have orginally been subspecies, fully capable of interbreeding. Eventually they attained full species status, and over time became the orders we know now. There were probably other lineages that didn't survive to the present.

At present we don't know what these ancestral species were like, but they may have been sort of shorebirdy, as in Feduccia's (1999) “transitional shorebirds” hypothesis. Near relatives to all modern birds include taxa such as Ichthyornis and Iaceornis (see, e.g., Longrich et al., 2011). Interestingly, Longrich et al. found fossils of one such relative, which they designate “Ornithurine C”, on both sides of the K/Pg boundary (K/T boundary). Since no representatives of Neoaves were included in the analysis, we cannot say if this taxon was closer to Neoaves than to Galloanserae and Paaleognathae.

Further Thoughts on the Polytomy

We now see why the relationships within Neoaves have been so problematic. The Vegavis fossil (Clarke et al., 2005) tells us that the Palaeognathae and Galloanserae lineages separated from the other birds in the Cretaceous. Neoaves seems to owe its orgina to the consequences of the bolide that crashed into Chicxulub at the end of the Cretaceous (currently estimated at 66.04 million years ago, see Renne et al., 2013).

The resulting environmental devastation wiped out many species, genera, families, and even orders of living organisms. Even some of those that survived proved unable to thrive in the new world of the Paleogene.

The surviving birds rapidly diversified following the impact, creating problems such as incomplete lineage sorting at supra-ordinal levels within Neoaves. Untangling this has been a protracted process. It is now twenty-five years after Sibley and Ahlquist, and the problem has still not been completely solved. Jarvis et al. (2014) threw a huge amount of data at the problem, three orders of magnitude more than Hackett et al. (2008). In spite of all this, Suh (2016) informs us that we are still left with an intractable polytomy in Neoves. As I update this in late 2023, the situation remains unchanged.

Perhaps new methods and still more data will lead to resolution of the problem, but we shouldn't count on it.


The oldest major division among living birds is between the Palaeognathae and Neognathae, as named by Pycraft (1900). Although the ratites had long been considered related, Pycraft's completed the group by including the Tinamous, creating the Palaeognathae. This consists of the volant tinamous, the non-volant ratites (ostriches, rheas, emus, cassowaries, kiwis), and their extinct relatives, which include moas, elephant birds, and the Lithornithiformes (sometimes called false tinamous).

Traditionally, the Palaeognathae were divided into the flightless ratites and the volant tinamous. Some of the earlier generic evidence seemed to support this (e.g., Haddrath and Baker, 2001).

This simple picture is now known to be wrong. An avalanche of more recent studies, including Chojnowski et al. (2008), Hackett et al. (2008), Harshman et al. (2008), Phillips et al. (2010), Faircloth et al. (2012), Haddrath and Baker (2012), J.V. Smith et al. (2013), Grealy et al. (2017), Yonezawa et al. (2017), Cloutier et al. (2019), Urantówka et al. (2020), Simmons et al. (2022), and Takezaki (2023), have come to a very different conclusion. The ratites are not a natural grouping! The ratites are not monophyletic.

Moas, Elephant Birds, and Lithornithiformes

The problem is that some of the ratites are more closely related to the volant tinamous than they are to other ratites. In particular, the ostriches are no more closely related to most of the other ratites than they are to the tinamous.

Palaeognathae tree

The diagram to the right shows the current situation. It includes not only living birds, but some extinct taxa. By using ancient DNA, we have been able to properly place the extinct Elephant Birds and Moas on the tree.

Unfortunately, ancient DNA only works if the fossils are not too ancient, so the extinct Lithornithiformes are a problem. Comparison of bones suggests that the most likely possibilities are that they are sister to the tinamous (Johnston, 2011; Nesbitt and Clarke, 2016), or are sister to all of the surviving paleognaths (Worthy et al., 2017; Yonezawa et al., 2017). Both possibilities are illustrated on the accompanying diagram by including the Lithornithiformes twice.

In fact, the diagram may still oversimplify the situation with the Lithornithiformes. They may not be a monophyletic group. See Nesbitt and Clarke (2016) and Widrig and Field (2022) for more on the Lithornithiformes.

What about the Rheas?

It's clear enough that the ostriches are the basal among the extant Palaeognathae. Once we get beyond the ostriches, things are not as clear-cut.

The remaining Palaeognathae comprise three groups: (1) rheas, (2) tinamous (and the recently extinct moas), and (3) cassowaries, emus, kiwis and the recently extinct elephant birds. These groups all separated over a short period of time, near the beginning of the Paleogene. E.g., Yonezawa et al. (2017) estimate the three groups separated form one another over a period of 4 million years.

So which group separated first? Even though it has been extensively studied, the short time period makes it hard to answer this question. Many analyses (e.g., Harshman et al., 2008; Phillips et al., 2010; J.V. Smith et al., 2013) favor putting the rheas first.

However, when Haddrath and Baker (2012) considered retroposons, the balance shifted to the tinamaou/moa clade, and that is the topology we formerly followed here. It is also the topology found by Cloutier et al. (2019), who used both coalescent and concatenated methods to address the question. The coalescent methods found that group (2), the tinamous and moas separated first. In contrast, concatenation favored group (1), the rheas.

They attributed this discrepancy to incomplete lineage sorting. This is not surprising as the lineages all arose over a period of about 4 million years. Because coalescent methods are generally more accurate, I assumed the moas and tinamous separated first. TiF has followed this starting in December 2009 (version 2.52 of this file).

Well, it is time for a change! More recent analyses such as Urantówka et al. (2020), Simmons et al. (2022), and Takezaki (2023) favor putting the rheas first, and I have returned to that arrangement.

Flightlessness Common in Ratites

This topology suggests that flightlessness has evolved at least 6 times (!) in the Palaeognathae: in ostriches, rheas, moas, elephant birds, emus/cassowaries, and kiwis. The fact that the tinamous can fligh implies flightlessness evolved at least four times. This is true even the rhea ancestors rafted across the then-narrow Atlantic, rather than flying. They still must have been capable of flight to pass it on to the tinamous. The distribution of the emus/cassowaries, elephant birds, kiwis suggests they each had volant ancestors, accounting for the other two times.

The chronogram in Haddrath and Baker (2012) suggests both the split of the rheas and Austalasian ratites and the split between the kiwis and emus/cassowaries occurred about 80 million years ago (mya). This timing is consistent with the kiwis originating due to the separation of Australia and New Zealand and with the ancestral rheas walking their way across the then-temperate Antarctica to South America. However, based on other chronograms such as Yonezawa et al., 2017, I think it likely that the rhea split occurred after the end of the Cretaceous, and the emu/cassowary clade branching off even later. (That's not the timing that they actually show, but I think it likely the rhea split was a bit later, after the Chicxulub meteor hit.)

Paleognath Geography

There is further relevant DNA evidence indicating that the extinct elephant birds of Madagascar are the closest relatives of the kiwis (Mitchell et al., 2014b), with a common ancestor perhaps 50 mya. This is a problem for vicariance theories driven by continental drift because Madagascar likely separated from Gondwana considerably earlier, perhaps 120 mya. In that case, the ancestor of the elephant birds would have had to fly, and we can infer that flight was indpendently lost in the rheas, emu/cassorwary, elephant bird, and kiwi lineages, a total of 6 losses of flight. This is not an unreasonable number. One only has to consider all of the flightless rails to see that.

Paleognath Orders

The Palaeognathae are divided into several orders in recognition of the great antiquity of the various branches, with the ostriches possibly dating back to the Cretaceous period. Phillips et al. (2010) estimate the ostriches diverged from the rest of the Palaeognathae about 60-95 mya, while Chojnowski et al. (2008) date this divergence around 64±22 mya, near the K/T (=K/Pg) boundary. Haddrath and Baker (2012) put it at 73-119 mya. Finally, mapping Jarvis et al. (2014) onto the phylogeny used here suggests that the split was about 84 mya (interval 58-96 mya).

The Cassowaries and Emus are much more closely related than the other paleognath groups, and so are placed in a single family: Casuariidae. Although the Moas and Elephant Birds are not part of the main TiF list, a genus-level phylogeny of both is given in the tree diagram. The Moa tree is based on Bunce et al. (2009).

Based on calibrated trees, I suspect that only the ostriches and possibly false tinamous survived the Chicxulub meteorite that ended Cretaceous and the non-avian dinosaurs.


Struthionidae: Ostriches Vigors, 1825

The authors for family-group names are mostly based on Bock (1994), while the authors for order-group names are primarily based on the series by Brodkorb (1963-1978). In both cases, additional sources have been consulted (e.g., Livezey and Zusi (2007)). In a number of cases, I have managed to examine the original (thank you Google Books!). The ICZN does not regulate order-group names. However, for parvorders (-ida), infraorders (-ides), suborders (-i), orders (-iformes), and superorders (-imorphae), I am attempting to follow similar rules. In particular, they are based on priority and the orginal use must been at an ordinal level and must be based on an included genus name (type genus) actually used by that author. The endings have been adjusted to modern usage.

1 genus, 2 species HBW-1

RHEIFORMES Forbes, 1884

Rheidae: Rheas Bonaparte, 1849

1 genus, 2 species HBW-1


The Dromaiidae (Emus) have been merged into the Casuariidae (cassowaries) because the split between them seems fairly recent. The molecular dates in Phillips et al. (2010) and in Haddrath and Baker (2012) already suggested they were so closely related that they could be treated as a single family. More precise dating by Prum et al. (2015) placed the split at about 10 million years ago, too close to even rank them as subfamilies.

Casuariidae: Emus and Cassowaries Kaup, 1847

2 genera, 4 species HBW-1

Heupink et al. (2011) argue that the extinct King Island Emu, Dromaius ater, was quite closely related to the extinct Tasmanian subspecies of the Emu, and is best considered a dwarf subspecies of Dromaius novaehollandiae. The Kangaroo Island Emu, D. baudinianus, seems likely to have been no more different from the Emu than the King Island Emu was, so I am also considering it a subspecies of the Emu, D. novaehollandiae.


Apterygidae tree

The order of the Kiwis reflects the phylogeny in Burbidge et al. (2003) and Shepherd et al. (2012). Burbidge et al. also provide evidence for recognizing 3 extant species of brown kiwi, as is done here. Shepherd et al. found evidence of an extinct northern clade of Little Spotted Kiwi that may deserve recognition as a separate species. The DNA samples of this potential species are from bones of unknown age.

Apterygidae: Kiwis G.R. Gray, 1840

1 genus, 5 species HBW-1


Tinamidae tree
Click for Tinamou species tree

The Tinamiformes (and their extinct Moa cousins) form the remaining branch of the Palaeognathae. The taxonomy of the Tinamous is now based on Musher et al. (2024) and SACC.

The age of the Tinamidae crown group is rather uncertain. Almeida et al. (Figure 3) give an age of 40 million years based on the complete data (black lines in their Fig. 3), but only about 30 million years based on the RAG2 gene (gray lines). To add to the confusion, Prum et al. (2015) puts it near 26 million years ago. Generally speaking, I prefer to go with Prum's number. As a result, the age estimates for genera must even younger than shown by the gray lines. In particular, Prum et al. place the basal split in Eudromiinae at about 21 mya and the split between Cryptura and Tinamus at about 17 mya. This compares with about 25 and 21 mya according to the gray lines.

The Tinamidae naturally divide into two subfamilies: Eudromiinae (Steppe Tinamous) and Tinaminae (Forest Tinamous) and 8 genera.

The current arrangement of the Tinamidae follows Figure 2 of Musher et al. (2024). The current species tree marks the species with new DNA data using bright red asterisks. Only one species was not sampled, the Slaty-breasted Tinamou, Tinamus boucardi. Its placement reflects hybridization in Honduras with the Thicket Tinamou, Tinamus cinnamomeus (Monroe, 1968).

Besides reordering the tinamous, the Musher et al. tree suggests that the Andean Tinamou, Nothoprocta pentlandii, represents two non-sister species. In Birds of the High Andes, Fjeldså and Krabbe (1990) noted that the Andean Tinamou consists of a brownish group of subspecies and a grayish group. Accordingly, I've split the Andean Tinamou, Nothoprocta pentlandii, into two species:

The New Tinamus, plus Cryptura and Crypturus

You may be puzzled by the way I use Tinamus. Wasn't Tinamus major the type? How can it be booted out of Tinamus? What happened?

Well, although people have treated Tinamus major as the type, it isn't! I don't know the complete story, but the genus Tinamus had been attributed to Latham (1790) until Hermann's 1783 Tabula affinitatum animalium was eventually noticed. This clearly has priority over Latham.

The problem is that the type must be among the species named in the original description. Hermann only mentioned soui. He also referred to a description of tinamous in Buffon, who included major and a few others, but did not explicitly refer to the species included in Buffon. Names included only by reference cannot be among the originally included nominal species. The ICZN rules are quite clear on this (Art. 67.2.3). They are also clear that the type must be an originally included nominal species. That means that Tinamus soui is the only possible type species for Tinamus.

This problem was pointed out in BirdForum by Laurent Raty. Some of the details are also mentioned on the Richmond cards for Tinamus here and here, where the reference to Buffon is accepted. And yes, Buffon is the same Buffon as in Buffon's needle problem. Using the (now incorrect) inclusion of Buffon's species, Apstein fixed the type as major in 1915. I'm not sure when the code explicitly barred inclusion by reference.

No one seems willing to follow the Code here, so I thought I would stir the pot by doing so. One problem with the Code is that it can cause (scientific) naming chaos. Ideally, the ICZN would decide in favor of stability here and elsewhere (e.g., Sulidae) and preserve traditional usage.

As major is also the type of Cryptura, I've put what had been the Tinamus species in Cryptura. What had been called Crypturellus is now mostly called Tinamus, with the others (cinereus and berlepschi) in Crypturus. Yes, it's confusing, but it's long past time to correct this.

Type Species

The table below lists the type species for each of the genera used by the current TiF list. Type species are noted on the species tree by leading black 5-pointed stars.

Genus Type Species Author Year

Tinamotis pentlandii Vigors 1837
Eudromia elegans I. Geoffroy Saint-Hilaire 1832
Nothura boraquira Wagler 1827
Rhynchotus rufescens von Spix 1825
Nothoprocta perdicaria Sclater & Salvin 1873
Nothocercus julius Bonaparte 1856
Crypturus cinereus Illiger 1811
Cryptura majora Vieillot 1816
Tinamus soui Hermann 1783

Compared to the earlier TiF version of the Tinamous (versions 3.03, March 18, 2018 and earlier) which had the same genera as version 13.2 of the IOC list, Taoniscus has been absorbed by Nothura, Tinamus is renamed Cryptura, Berlepsch's Tinamou, Crypturellus berlepschi and Cinereous Tinamou, Crypturellus cinereus have been moved to Crypturus, and the remainder of Crypturellus is now called Tinamus.

What about the Chaco Nothrua?

Hayes et al. (2018) showed that there is no reason to consider the Chaco Nothura, Nothura chacoensis, distinct from the Spotted Nothura, Nothura maculosa. As a result, I now consider the Chaco Nothura to be a subspecies of the Spotted Nothura, Nothura maculosa.

Tinamidae: Tinamous G.R. Gray, 1840

9 genera, 47 species HBW-1

Eudromiinae: Steppe Tinamous Bonaparte 1854

  1. Puna Tinamou, Tinamotis pentlandii
  2. Patagonian Tinamou, Tinamotis ingoufi
  3. Quebracho Crested-Tinamou, Eudromia formosa
  4. Elegant Crested-Tinamou, Eudromia elegans
  5. White-bellied Nothura, Nothura boraquira
  6. Dwarf Tinamou, Nothura nanus
  7. Lesser Nothura, Nothura minor
  8. Darwin's Nothura, Nothura darwinii
  9. Spotted Nothura, Nothura maculosus
  10. Brushland Tinamou, Rhynchotus cinerascens
  11. Red-winged Tinamou, Rhynchotus rufescens
  12. Huayco Tinamou, Rhynchotus maculicollis
  13. Curve-billed Tinamou, Nothoprocta curvirostris
  14. Ornate Tinamou, Nothoprocta ornata
  15. Brown Andean Tinamou, Nothoprocta oustaleti
  16. Taczanowski's Tinamou, Nothoprocta taczanowskii
  17. Gray Andean Tinamou, Nothoprocta pentlandii
  18. Chilean Tinamou, Nothoprocta perdicaria

Tinaminae: Forest Tinamous G.R. Gray, 1840

  1. Tawny-breasted Tinamou, Nothocercus julius
  2. Highland Tinamou, Nothocercus bonapartei
  3. Hooded Tinamou, Nothocercus nigrocapillus
  4. Great Tinamou, Cryptura majora
  5. Gray Tinamou, Cryptura tao
  6. Solitary Tinamou, Cryptura solitaria
  7. White-throated Tinamou, Cryptura guttata
  8. Black Tinamou, Cryptura osgoodi
  9. Berlepsch's Tinamou, Crypturus berlepschi
  10. Cinereous Tinamou, Crypturus cinereus
  11. Tepui Tinamou, Crypturus ptaritepui
  12. Brown Tinamou, Tinamus obsoletus
  13. Tataupa Tinamou, Tinamus tataupa
  14. Small-billed Tinamou, Tinamus parvirostris
  15. Variegated Tinamou, Tinamus variegatus
  16. Little Tinamou, Tinamus soui
  17. Bartlett's Tinamou, Tinamus bartletti
  18. Barred Tinamou, Tinamus casiquiare
  19. Rusty Tinamou, Tinamus brevirostris
  20. Undulated Tinamou, Tinamus undulatus
  21. Yellow-legged Tinamou, Tinamus noctivagus
  22. Black-capped Tinamou, Tinamus atrocapillus
  23. Pale-browed Tinamou, Tinamus transfasciatus
  24. Thicket Tinamou, Tinamus cinnamomeus
  25. Slaty-breasted Tinamou, Tinamus boucardi
  26. Choco Tinamou, Tinamus kerriae
  27. Red-legged Tinamou, Tinamus erythropus
  28. Gray-legged Tinamou, Tinamus duidae
  29. Brazilian Tinamou, Tinamus strigulosus

Previous Page Next Page