CHARADRIIFORMES Huxley, 1867
The Charadriiformes include 19 families, containing over 120 genera and close to 400 species, ranging from shorebirds to alcids, terns and gulls. The Charadriiformes have been carefully studied in recent years and DNA methods have proven especially effective at unraveling the taxonomy. We not only know how most of the various families relate, but we also have a good handle on many of the genera (the large white-headed gulls (genus Larus) continue to puzzle).
There is a lot of evidence for monophyly of the Charadriiformes as constituted here (e.g., Ericson et al., 2003a; Paton et al., 2003; Cracraft et al., 2004; Thomas et al., 2004a; Paton and Baker, 2006; Baker et al., 2007; Fain and Houde, 2007, Černý and Natale, 2021). Many studies have found that gulls and alcids are closely related to the shorebirds. Collectively, these analyses have made it quite clear that the sandgrouse (Pteroclidae) and bustards (Otididae) are not Charadriiformes. They have also shown that the buttonquail (Turnicidae) and Plains-wanderer (Pedionomidae) are Charadriiformes. The position of the buttonquail is also supported by morphology (Mayr, 2008a).
The last family to join the Charadriiformes was the buttonquails (Turnicidae). At one point, they were considered related to quail, and they were placed in the Galliformes (e.g., Gadow, 1892). However, they are actually quite different from and were later grouped with the cranes and rails (Gruiformes). Sibley, Ahlquist, and Monroe (1988) listed them in their own separate group. van Tuinen, Sibley, and Hedges (2000) had Turnix sister to Larus, but not Charadriius, which was also nearby. Paton et al. (2003), in their study of Charadriiformes, attempted to use Turnix as an outgroup. Imagine their surprise when they found it was nested within Charadriiformes. Indeed, the found it was sister to what we call suborder Lari, as it is today. At this point, there are no plausible additions to the Charadriiformes. They have all be tested genetically, and the Turnicidae were the last.
The basic structure of this order is now pretty well worked out. However, there are still some details missing. Sequencing remains spotty for some genera. Further, there is still some uncertainty regarding the precise position of certain genera, and species within them. For the latter, the Herring Gull complex comes to mind.
To help make the taxonomy clear, the Charadriiformes have been divided into 4 suborders: Charadrii, Limicoli, Turnici, and Lari. The tree below shows how it all fits together, including the division into parvorders (-ida) and superfamilies (-oidea), ending with the 20 shorebird families (-idae). The 3-page genus-level Charadriiformes tree, leaves off the parvorders and superfamilies to focus on families, genera, and species. It also includes subfamilies (-inae) and tribes (-ini).
|Click for Charadriiformes|
family and genus tree
Another problem that cropped up around the turn of the century was the fact that standard molecular clocks performed poorly in large-scale genetic analyses. That lead to the use of calibration. One early calibration effort was van Tuinen and Hedges (2001). As is usual with calibration studies, their ages were too old rather than too young (see their Fig. 3). This has been repeatedly seen in calibration efforts, including that of Baker et al. (2007), who focused on the Charadriiformes.
Calibrations have been refined since, but there's still considerable uncertainty concerning the age of the shorebird clades and species. Moreover, in practice, calibration is based toward older ages. This can be overcome if you have enough good calibration fossils. So far we don't, but Černý and Natale (2021) made a good try. Moreover, they are the first to properly vet their calibration fossils. This is especially important when dealing with any paleontological literature that is older than about 2010. In the older literature, they often had strange ideas about the relation between fossils and modern taxa. Kudos to Černý and Natale for dealing with this problem. The DNA did not get a similar level of scrutiny in their unpublished draft, but they're working on correcting that (Černý, pers. comm.).
I think even Černý and Natale (2021) still have many dates that are too old. Many of these occur in branches far from calibration points. Where there are nearby calibration points, the dating is considerably impoved compared with other calibrated studies of the Charadriiformes. Compare Paton et al. (2003) or Baker et al. (2007). One clue that there are still some problems involves the prolonged lack of speciation in much of the Černý and Natale tree (look at the tips). Another common artifact in calibration studies is a regular proportional spacing out of extra time.
Černý and Natale's (2021) use of many more calibration points have had an beneficial effect and some parts of the tree are starting to look pretty reasonable. I'm using their age data with care when designating higher-level taxa, and also when deciding whether to use additional genera (I generally consider shorebird families over-lumped at the genus level).
Charadrii Huxley, 1867
Chionida Sharpe, 1890
The Burhinidae and Chionidae are more closely related to each other than to the other shorebirds.
Chionidae: Sheathbills Lesson, 1828
2 genera, 3 species HBW-3
It has become increasingly clear that the Magellanic Plover is fairly closely related to the Sheathbills. Černý and Natale (2021) estimated their common ancestor lived within the last 20 million years. As a result, I'm treating both groups as subfamilies of a single family (the name Chionidae has priority---1828 vs. 1975).
Pluvianellinae: Magellanic Plover Jehl, 1975
- Magellanic Plover, Pluvianellus socialis
Chioninae: Sheathbills Lesson, 1828
- Snowy Sheathbill, Chionis albus
- Black-faced Sheathbill, Chionis minor
Burhinidae: Thick-knees Mathews, 1912 (1840)
4 genera, 10 species HBW-3
Before using Černý and Natale (2021) on the thick-knees, we have to answer the key question: What is Burhinus magnirostris? That name has been used to refer to both the Beach Stone-Curlew and the Bush Stone-Curlew. So why do two birds have the same scientific name, and which one is meant here.
The Two Birds Called Burhinus magnirostris
Although the Beach Stone-Curlew, often called Esacus magnirostris, and the Bush Stone-Curlew, sometimes called Burhinus magnirostris are two entirely different birds, they are both magnirostris. Here are the original forms of these names:
- Beach Stone-Curlew: Oedicnemus magnirostris Vieillot 1818
- Bush Stone-Curlew: Charadrius magnirostris Latham 1801
So what's going on here, and why is the Bush Stone-Curlew often called Burhinus grallarius? It all has to do with rules governing priority of names, the ICZN Code, overseen by the International Commission on Zoological Nomenclature.
We'll start with the easy one first. When the Bush Stone-Curlew as originally named by Latham, he gave it three names, the other two being grallarius and frenatus. Since these were used in the same publication, none was older, which would have established priority. In a case like this, someone has to pick. Well, evidently no one did, at least explicitly. No one pointed to the three names and said “this one”!
So magnirostris came into use for the Bush Stone-Curlew. Finally, Gould (1845) picked grallarius. This took a long time to come into regular use, but has been followed by much of the recent literature. Nonetheless, one sometimes still sees Burhinus magnirostris for the Bush Stone-Curlew.
One might think that this would allow us to use magnirostris for the Beach Stone-Curlew. However, if both are placed in the genus Burhinus, and they sometimes are, Latham's name still has priority over Vieillot's even though it is not in use. That means that Burhinus magnirostris cannot be used for the Beach Stone-Curlew, although it sometimes is, even in the 21st century.
Technically, Vieillot's name is a secondary junior homonym of Latham's. It's junior because Latham got there first, and secondary because Latham's name is not actually in use (officially). Nonetheless, that is enough to prevent it from being used when both are in the same genus.
Under the current rules, that would mean that magnirostris would be available for the Beach Stone-Curlew provided that it was not in the same genus as the Bush Stone-Curlew. However, the rule used to be different, and in the interest of maintaining the same name when possible, decisions made under the previous rules are still binding. The old rule holds prior to 1961, and would have prohibited use of magnirostris for the Beach Stone-Curlew under any circumstances. This holds provided someone published a paper that pointed it out. Just as someone (eventually Gould) had to choose the right name for the Bush Stone-Curlew, someone needed to address the problem.
That brings us to Meinertzhagen (1924), well before the cutoff of 1961. She put all of the Burhinidae in a single genus. In spite of Gould (1845), magnirostris was still being used for both species and Meinertzhagen had to do something about it. In her species account for the Beach Stone-Curlew (Meinertzhagen, 1924) Meinertzhagen rejected Wagler's name giganteus as unidentifiable (pg. 351). She also rejected the name major). Next in line was neglectus Matthews, 1912. Meinertzhagen was able to examine the type specimen, and verify its identity, and the Beach Stone-Curlew became Burhinus neglectus. Since this happened before 1961, Article 59.3 of the Code applies. This makes the application of magnirostris to the Beach Stone-Curlew permanently invalid. Unfortunately, it doesn't stop people from using it and causing confusion.
An internet search reveals that the name has gotten some use, but the Beach Stone-Curlew is often placed in Esacus and called Esacus magnirostris, contrary to Article 59.3. One example is Hayman et al. (1986), which uses magnirostris for both Bush and Beach Stone-Curlews.
Relatively recent discussions of this issue can be found in Christidis and Boles (1994, 2008) and Hume's (1996) article on Burhinidae in HBW-3 (pg. 350) as well as a recent thread on BirdForum. That's why you see Beach Stone-Curlew named Esacus neglectus in the list below. Hope I have it all straight!
That brings us back to Černý and Natale (2021).
What is Burhinus magnirostris?
So what do Černý and Natale (2021) mean when they say Burhinus magnirostris? Figure A.2 shows they do not use any genetic information from it. So the only information is morphological, from Strauch (1978). According to Černý it is the Beach Stone-Curlew, called Esacus magnirostris by Strauch that they mean by Burhinus magnirostris. He noted they also used morphological data for the Bush Stone-Curlew, called Burhinus magnirostris by Strauch, which we call Burhinus grallarius, and that they were careful to keep them straight. That makes it clear that Černý and Natale intend that Burhinus magnirostris refer to the Beach Stone-Curlew and that Burhinus grallarius refer to the Bush Stone-Curlew. You'd think that would resolve the problem.
It dosen't because they also used sequences from GenBank from what GenBank identifies as Bush Thick-knee, Burhinus grallarius (aka Bush Stone-Curlew). If we look in GenBank, we find a number of genes listed as Burhinus grallarius, the Bush Thick-knee. These genes were sequenced variously for the papers by Paton et al. (2003), Paton and Baker (2006), and Baker et al. (2012). They ultimately sampled three specimens from the Royal Ontario Museum (Toronto). But the papers don't mention obtaining any genes from specimens of the Bush Thick-knee. The papers tell us the birds involved were Burhinus magnirostris (!) and helpfully clarify its identity by giving the English name, Beach Thick-knee (i.e., Beach Stone-Curlew).
All of the DNA they extracted refers to vouchered specimens AJB6174, AJB6175, and AJB6177 (AJB = Allan J. Baker). In fact, the only real Bush Thick-knee genes represented in GenBank seem to be microsatellites. There's one other submission that claims to be cyt-b from a Bush Thick-knee. However, that DNA sample is from blood taken from a mosquito that bit a bird. The bird was identified by noticing the partial cty-b sample was identical to a portion of the cyt-b sample labelled voucher AJB6177, and sampled by Paton and Baker (2006). In other words, it's really a Beach Stone-Curlew too.
So the answer is that in Burhinus magnirostris refers to the morphological Beach Stone-Curlew, while Burhinus grallarius refers to a chimera consisting of DNA from the Beach Stone-Curlew, with the morphology of a Bush Stone-Curlew. All these difficulties stem from using the same name for two birds.
The DNA barcoding database BOLD seems to include both Beach and Bush Stone-Curlews. See Laurent Raty's discussion on BirdForum.
Buhinidae Taxonomic Notes
Now that we have probable ID for "Burhinus magnirostris" as the Beach Stone-Curlew, we can use Černý and Natale (2021), we to sort out the Burhinidae in a way consistent with the available genetic evidence.
Thick-knees: Based on Paton et al. (2003) and Černý and Natale, (2021), there seem to be four main clades in Burhinus. I will treat them all as separate genera.
- The new world clade includes the Double-striped Thick-knee, "Burhinus" bistriatus, and the Peruvian Thick-knee, "Burhinus" superciliaris. There is no available genus name, so I will call these "Burhinus" for now.
- The Bush Stone-Curlew, Burhinus grallarius. This is Burhinus, now reduced to a single species.
- The Great and Beach Stone-Curlews, still called genus Esacus.
- The remaining five species, formerly included in Burhinus, get the name Oedicnemus (Temminck 1815, type oedicnemus).
|Click for Burhinidae|
The relation between the other three clades is still unresolved because the Bush Stone-Curlew remains unsampled. It is believed to be close to Esacus, so I treat the three clades as a trichomoty. I restrict Burhinus to a single species, the Bush Stone-Curlew, Burhinus grallarius. A second clade is Esacus, containing the Beach and Great Stone-Curlews. Finally, the third clade contains the other species. It takes the genus name Oedicnemus (Temminck 1815, type oedicnemus).
Although Černý and Natale (2021) had no DNA from either Esacus, Paton et al. (2003) sequenced the RAG-1 gene for the Beach Stone-Curlew, Esacus magnirostris, and several other thick-knees. The sample (AY228769) is available from GenBank, but its name was erroneously corrected to Burhinus grallarius. How did that happen? It's simple. There are two thick-knees that use the name magnirostris, a recipe for confusion.
- Double-striped Thick-knee, "Burhinus" bistriatus
- Peruvian Thick-knee, "Burhinus" superciliaris
- Bush Stone-Curlew, Burhinus grallarius
- Great Stone-Curlew, Esacus recurvirostris
- Beach Stone-Curlew, Esacus neglectus
- Spotted Thick-knee, Oedicnemus capensis
- Water Thick-knee, Oedicnemus vermiculatus
- Eurasian Stone-Curlew, Oedicnemus oedicnemus
- Senegal Thick-knee, Oedicnemus senegalensis
- Indian Stone-Curlew, Oedicnemus indicus
Pluvianida: Egyptian Plover Informal?
The Egyptian Plover or Crocodile-bird is sister to all of the remaining Charadrii.
Pluvianidae: Egyptian Plover Reichenbach, 1848
1 genus, 1 species Not HBW Family
Although it is sometimes put in its own family, the Egyptian Plover is typically considered a member of the Glareolidae (pratincoles and coursers). Ericson et al. (2003a), Baker et al. (2007), and Fain and Houde (2007) make clear it is nowhere close to the Glareolidae. All three found it to be basal in the Charadrii (in our sense). Further, all found it to be sister to the remaining Charadrii, which justifies placing it in its own family, and even its own parvorder (Pluvianida).
- Egyptian Plover / Crocodile-bird, Pluvianus aegyptius
Charadriida Huxley, 1867
The Charadriida consist of two clades, which I rank as superfamilies: Recurvirostroidea and Charadrioidea.
Recurvirostroidea Bonaparte, 1831
The superfamily Recurvirostroidea contains three families: Ibidorhynchidae (Ibisbill), Recurvirostridae (Stilts and Avocets), and Haematopodidae (Oystercatchers).
Ibidorhynchidae: Ibisbill Bonaparte 1856
1 genus, 1 species HBW-3
The Ibisbill is the basal branch in Recurvirostroidea.
- Ibisbill, Ibidorhyncha struthersii
Recurvirostridae: Stilts, Avocets Bonaparte, 1831
3 genera, 9 species HBW-3
|Click for Recurvirostridae|
The current arrangement is based on Černý and Natale, (2021). The Banded Stilt, Cladorhynchus leucocephalus, is basal in family Recurvirostridae, which has been rearranged accordingly.
I leave Recurvirostridae and Haematopodidae as separate families, but further reductions in their ages could lead to them being combined in a single family.
Baker et al. (2007) gave a different arrangement of the stilt and avocet genera. However, this seems to be driven by an ND2 sequence for Recurvirostra americana that appears to belong to a Painted-snipe. See Raty's comments on BirdForum for more details.
- Banded Stilt, Cladorhynchus leucocephalus
- Black-winged Stilt, Himantopus himantopus
- White-headed Stilt, Himantopus leucocephalus
- Black-necked Stilt, Himantopus mexicanus
- Black Stilt, Himantopus novaezelandiae
- Pied Avocet, Recurvirostra avosetta
- American Avocet, Recurvirostra americana
- Red-necked Avocet, Recurvirostra novaehollandiae
- Andean Avocet, Recurvirostra andina
Haematopodidae: Oystercatchers Bonaparte, 1838
2 genera, 13 species HBW-3
|Click for Haematopodidae|
The arrangement of the oystercatchers here is based on not yet published work by Senfeld et al. (2020b). You'll notice that it differs quite a bit from Černý and Natale (2021). However, if you focus on the species where Černý and Natale relied on sequence data (red stars on the tree), both agree. The differences are because Černý and Natale (2021) used morphological data to place the other species, and it was misleading, as morphological data sometimes is. In fact, that problem is part of why TiF was born, to present the massive taxonomic revisions based on genetic sequencing.
There's a deep divide between the Old World and New World Oystercatchers, so I have placed the New World species in the genus Prohaematopus (Matthews 1913, type ater).
Far Eastern Oystercatcher, Haematopus osculans: Based on Senfeld et al. (2020b), the Far Eastern Oystercatcher, Haematopus osculans, has been split from the Eurasian Oystercatcher, Haematopus ostralegus.
In spite of Senfeld et al.'s (2020a) recommendation to merge it into the Eurasian Oystercatcher, Haematopus ostralegus, I continue to list the extinct Canary Islands Oystercatcher, Haematopus meadewaldoi, as a separate species. See Collar et al. (2021) for details. The fact that the taxa appear to have been acting as biological species makes this a very different case than the Redpolls.
- Magellanic Oystercatcher, Prohaematopus leucopodus
- Blackish Oystercatcher, Prohaematopus ater
- Black Oystercatcher, Prohaematopus bachmani
- American Oystercatcher, Prohaematopus palliatus
- Sooty Oystercatcher, Haematopus fuliginosus
- African Oystercatcher, Haematopus moquini
- Eurasian Oystercatcher, Haematopus ostralegus
- †Canary Islands Oystercatcher, Haematopus meadewaldoi
- Far Eastern Oystercatcher, Haematopus osculans
- Pied Oystercatcher, Haematopus longirostris
- Chatham Oystercatcher, Haematopus chathamensis
- Variable Oystercatcher, Haematopus unicolor
- South Island Oystercatcher, Haematopus finschi
Charadrioidea: Plovers Leach, 1820
There are two families in the superfamily Charadrioidea: Pluvialidae (golden-plovers) and Charadriidae (plovers and dotterels).
Pluvialidae: Golden-Plovers MacGillivray, 1852
1 genus, 4 species Not HBW Family
One big surprise to come out of the molecular data is that the golden-plovers (and Black-bellied Plover) are not that closely related to the rest of the plovers. Several papers have even suggested that they are actually closer to the stilts, avocets, oystercatchers, and ibisbill (Ericson et al., 2003a; Baker et al., 2007; Fain and Houde, 2007).
Additional DNA, with more genes and more taxa, has clarified matters. The more radical suggestions are most likely incorrect, with the golden-plovers sister to the other plovers (Charadriidae, not the Magellanic or Egyptian Plovers). However, they are on a deep branch. Černý and Natale (2021) estimate their common ancestor lived roughly 39 mya. I think its somewhat less than that, but still sufficiently long ago to treat them as a separate family from Charadriidae.
- Black-bellied Plover / Gray Plover (Pluvialis squatarola)
- American Golden-Plover (Pluvialis dominica)
- European Golden-Plover (Pluvialis apricaria)
- Pacific Golden-Plover (Pluvialis fulva)
Charadriidae: Plovers, Dotterels Leach, 1820
22 genera, 63 species HBW-3
|Click for Charadriidae|
The last two editions of the Howard and Moore Checklist (Dickinson, 2003; Dickinson and Remsen, 2013) divided the plovers into three subfamilies: Pluvialinae, Vanellinae, and Charadriinae. Moreover, most of the species were in two genera: Vanellus and Charadrius. The organization is rather different here. Pluvialinae has been promoted to its own family. Problems with that classification appeared already in Christian et al. (1992). These problems intensified with Joseph et al. (1999). Both demonstrated that Charadriinae and Charadrius were natural groups. In retrospect, we can even see the current TiF organization in Joseph et al.
The latest TiF reoganization of the plovers draws heavily on Figure A.5 from Černý and Natale (2021), with the assistance of Barth et al. (2013) and Dos Remedios et al. (2015). I have selected Figure A.5, the RAxML analysis for the Charadriidae because it has better support in the Charadriidae than the ExaBayes analysis of Figure A.7.
There are a couple of issues with the DNA used by Černý and Natale (2021). One involves the Long-billed Plover, Thinornis placidus, where a Kentish Plover gene has been mixed in. See Päckert (2021) and Sangster and Luksenburg (2021) for more on this and other issues with published and archived genes.
Interestingly, Päckert's COI tree is also misleading for placidus, as a comparison with the 6 single-gene trees in Dos Remedios et al. (2015) shows. The placement of placidus sister to dubius in TiF tree is based on Dos Remedios et al. (2015).
The other problem involves one or both of the Caspian Plover, Eupoda asiatica, and the New Zealand Plover, Anarhynchus obscurus. Černý and Natale (2021) give three phylogenies based purely on genetics: ASTRAL-III, RAxML, and ExaBayes. These two species are sister in two of those, RAxML and ExaBayes, while the ASTRAL-III tree is consistent with the TiF tree, and with previous results (Barth et al., 2013,, dos Remedios et al., 2015). I don't really know what went wrong there.
Dos Remedios et al. (2015) and Barth et al. (2013) agree on the Eupoda clade, which contains the Caspian Plover and that the Wrybill, frontalis, New Zealand Plover, obscurus, and Double-banded Plover, bicinctus form a clade. There is some disagreement on the branching order. I've given preference to Dos Remedios et al. because they use more data than Barth et al. In fact, frontalis and obscurus show up as sister species in many of Dos Remedios's gene trees. The HBW Illustrated Checklist (del Hoyo and Collar, 2014) recognizes the two subspecies of the New Zealand Plover as separate species.
The names cucullatus and rubricollis: One point to clarify before considering the phylogeny involves the Hooded Dotteral. If you look at the papers cited, you'll notice that some refer to the Hooded Dotteral as rubricollis rather than cucullatus. There are two competing names, Charadrius rubricollis Gmelin, 1789 and Charadrius cucullatus Vieillot, 1818. Obviously, rubricollis is older, so why is cucullatus used?
The answer is in Mcallan and Christidis (1998) and Olson (1998). The short version of the story is that Gmelin's name rubricollis is based on Latham's description of a bird he called the Red-necked Plover, an odd name for the Hooded Dotterel, which doesn't have a red neck. Latham's description seems to have been based on drawings by Ellis from Cook's third voyage, but apparently he conflated drawings of the Dotterel and a Red-necked Phalarope! Mcallan and Christidis. Mcallan and Christidis attempted to correct this, but failed to correctly follow the ICZN Code. Olson corrected their correction, putting rubricollis among the synonyms of the Red-necked Phalarope. This automatically makes cucullatus the name of the Hooded Dotterel.
Charadriidae à la TiF
One big change we see by basing the phylogeny on Černý and Natale (2021) is that we have four subfamilies rather than three. The Tawny-throated Dotterel, Oreopholus ruficollis is sister to the remaining plovers, and is placed in its own subfamily, Oreopholinae. The rest of the organization is similar to earlier editions of TiF. The next branch is the subfamily Charadriinae.
Within Charadriinae, the Rufous-chested Dotterel (or Plover) is sister to the Diademed Sandpiper-Plover. They are only distant relative, and I previously placed the Sandpiper-Plover in the monotypic genus Zonibyx (Reichenbach, 1852). These sister taxa are basal in Charadriinae and the Eurasian Dotterel, Eudromias morinellus is the next branch.
At this point we have some rearrangement compared with pre-2021 editions of TiF, as the Three-banded Plover, Afroxyechus tricollaris, is thought to be allied with the Thinornis group. The tree has two branches here, a narrow Charadrius consisting of four species, and a clade containing Thinornis. Livezey (2010) suggested separating the first three species in this clade as Afroxyechus, but they are not sister species. Rather, they form a grade. They also seem to be distantly enough related to each other and the Thinornis group that we treat them as genera. Unfortunately, I couldn't find available names for two of them, so they get temporary designations:
- The Hooded Dotterel is referred to as "Thinornis" cucullatus
- Forbes's Plover is referred to as "Afroxyechus" forbesi
- The Three-banded Plover retains the name Afroxyechus tricollaris. The genus Afroxyechus (Mathews, 1913) is monotypic.
The remaining plovers divide into two subfamilies, the lapwings (Vanellinae) and the remaining plovers (Anarhynchinae). Although nine of the 24 lapwings have been sequenced, the diversity of plumage and form among the lapwings makes it hard to construct a reliable phylogeny. They are so diverse that 80% are the type of some genus.
The lapwings start with the Pied Lapwing, cayanus. It's not entirely clear that it is actually in the lapwing subfamily. The ASTRAL-III tree suggested it is sister to Oreopholus, the RAxML tree separated them, but would put the Pied Lapwing in its own subfamily, between Oreopholus and Charadriinae, and the ExaBayes tree has it sister to a combined Vanellinae plus Anarhynchinae. Finally, the total evidence tree has it sister to the rest of the Vanellinae.
I'm not sure what to think and have gone with the last. The thing that is clear is that the next (or basal!) group in the Vanellinae consists of the Northern, Southern, and Andean Lapwings, and that group seems sister to the remaining lapwings. The division between the Northern Lapwing and the Southern and Andean Lapwings appears deep, so I use four genera for the lapwings:
- The Pied Lapwing / Pied Plover is transferred to the monotypic genus Hoploxypterus (Bonaparte, 1856)
- Vanellus is restricted to the Northern Lapwing
- Southern Lapwing and Andean Lapwing are transferred to Belonopterus (Reichenbach 1852, type chilensis)
- Until we know more, the other 20 lapwings have been placed in Hoplopterus (Bonaparte 1831, type spinosus). I did not find the total evidence tree useful here as it contradicts the available genetic data. I retained the previous arrangement, with the above genera pulled out as above.
The last subfamily is Anarhynchinae. Erythrogonys and Peltohyas are basal, in that order. The other species were all once in Charadrius. I treat them as five clades, and the divisions within one of those clades are deep, requiring a total of seven genera.
The first branch is the sand-plover clade, Eupoda (J.F. Brandt 1845, type asiatica). That is followed by a clade of three species I previously put in genus Anarhynchus, but the divisions within it seem relatively deep, so I've split it into three genera. That gives us
- The Double-banded Plover, Anarhynchus bicinctus, is transferred to the monotypic genus Nesoceryx (Mathews, 1920).
- The New Zealand Plover, Anarhynchus obscurus, is transferred to the monotypic genus Pluviorhynchus (Bonaparte 1856).
- The Wrybill remains Anarhynchus frontalis (Quoy and Gaimard, 1830).
I had previously moved the remaining plovers from Charadrius to Ochthodromus (Reichenbach 1852, type wilsonia). Although some uncertainty, there appear to be three clades in Ochthodromus. It seems appropriate to separate them as genera.
- Leucopolius (Bonaparte 1856, type marginatus) includes the Kentish Plover, Leucopolius alexandrinus, and close relatives
- Helenaegialus (Mathews 1913, type sanctaehelenae) includes thoracicus, pecuarius, and sanctaehelenae.
- Ochthodromus (Reichenbach 1852, type wilsonia). Besides Wilson's (wilsonia) and Collared Plovers (collaris), it includes the Mountain Plover (montanus) and Puna (alticola) and Two-banded Plovers (falklandicus).
There's some possibility that Ochthodromus could be further split into three groups.
The taxonomic status of the Kentish-Snowy Plover complex has been controversial. Recent work by Küpper et al. (2009) found that the Kentish, Snowy, and White-fronted Plovers represented independent groups, with little or no evidence of gene flow between them. Not only does this separation appear to be long-standing, but the White-fronted Plover seems to be more closely related to the Kentish Plover than to the Snowy Plover. Accordingly, the Kentish (L. alexandrinus) and Snowy (L. nivosus) Plovers are treated as separate species below.
Rheindt et al. (2011b) provide further evidence in favor of separating Kentish and Snowy Plovers. They also found that the Malaysian Plover, Leucopolius peronii, is member of the group. It is most closely related to the White-fronted Plover, Leucopolius marginatus. Both are more closely related to the Kentish Plover than to the Snowy Plover.
The White-faced Plover, L. alexandrinus dealbatus, differs somewhat from the Kentish Plover in plumage, size, and ecologically (Bakewell and Kennerly, 2008; Kennerly et al., 2008). Whether it is a separate species remains a question. The HB2 Illustrated Checklist (del Hoyo and Collar, 2014) recognizes it as a separate species, but Rheindt et al. (2011b) found no genetic evidence supporting this.
- Tawny-throated Dotterel, Oreopholus ruficollis
Charadriinae Leach, 1820
- Diademed Sandpiper-Plover, Phegornis mitchellii
- Rufous-chested Dotterel / Rufous-chested Plover, Zonibyx modestus
- Eurasian Dotterel, Eudromias morinellus
- Killdeer, Charadrius vociferus
- Common Ringed Plover, Charadrius hiaticula
- Semipalmated Plover, Charadrius semipalmatus
- Piping Plover, Charadrius melodus
- Hooded Dotterel, "Thinornis" cucullatus
- Forbes's Plover, "Afroxyechus" forbesi
- Three-banded Plover, Afroxyechus tricollaris
- Little Ringed Plover, Thinornis dubius
- Long-billed Plover, Thinornis placidus
- Black-fronted Dotterel, Thinornis melanops
- Shore Dotterel, Thinornis novaeseelandiae
Vanellinae: Lapwings Bonaparte, 1842
- Pied Lapwing / Pied Plover, Hoploxypterus cayanus
- Northern Lapwing, Vanellus vanellus
- Southern Lapwing, Belonopterus chilensis
- Andean Lapwing, Belonopterus resplendens
- Long-toed Lapwing, Hoplopterus crassirostris
- Blacksmith Lapwing, Hoplopterus armatus
- Spur-winged Lapwing, Hoplopterus spinosus
- River Lapwing, Hoplopterus duvaucelii
- Black-headed Lapwing, Hoplopterus tectus
- Yellow-wattled Lapwing, Hoplopterus malarbaricus
- White-crowned Lapwing, Hoplopterus albiceps
- Senegal Lapwing, Hoplopterus lugubris
- Black-winged Lapwing, Hoplopterus melanopterus
- Crowned Plover / Crowned Lapwing, Hoplopterus coronatus
- African Wattled Lapwing, Hoplopterus senegallus
- Spot-breasted Lapwing, Hoplopterus melanocephalus
- Brown-chested Lapwing, Hoplopterus superciliosus
- Gray-headed Lapwing, Hoplopterus cinereus
- Red-wattled Lapwing, Hoplopterus indicus
- †Javan Lapwing, Hoplopterus macropterus
- Banded Lapwing, Hoplopterus tricolor
- Masked Lapwing, Hoplopterus miles
- Sociable Lapwing, Hoplopterus gregarius
- White-tailed Lapwing, Hoplopterus leucurus
Anarhynchinae Baird, Brewer, and Ridgway, 1884
- Red-kneed Dotterel, Erythrogonys cinctus
- Inland Dotterel, Peltohyas australis
- Caspian Plover, Eupoda asiatica
- Oriental Plover, Eupoda veredus
- Greater Sand-Plover, Eupoda leschenaultii
- Lesser Sand-Plover, Eupoda mongola
- Double-banded Plover, Nesoceryx bicinctus
- New Zealand Plover, Pluviorhynchus obscurus
- Wrybill, Anarhynchus frontalis
- Red-capped Plover, Leucopolius ruficapillus
- Snowy Plover, Leucopolius nivosus
- Chestnut-banded Plover, Leucopolius pallidus
- Kentish Plover, Leucopolius alexandrinus
- Javan Plover, Leucopolius javanicus
- White-fronted Plover, Leucopolius marginatus
- Malaysian Plover, Leucopolius peronii
- Madagascan Plover, Helenaegialus thoracicus
- Kittlitz's Plover, Helenaegialus pecuarius
- St. Helena Plover, Helenaegialus sanctaehelenae
- Mountain Plover, Ochthodromus montanus
- Wilson's Plover, Ochthodromus wilsonia
- Collared Plover, Ochthodromus collaris
- Puna Plover, Ochthodromus alticola
- Two-banded Plover, Ochthodromus falklandicus
Limicoli Garrod, 1873
The term Scolopaci (Strauch 1978?) seems to be the current fashion. However, Limicoli has a lot of priority and a long history of use (often as Limicolae).
Parrida Sharpe, 1891
The name was introduced by Sharpe (1891) as suborder Parrae in a paper on classification. If you wonder why he called it Parrae, it is enough to realize that Linnaeus called the Wattled Jacana Parra jacana. You also see it in the jacana genus names Microparra and Irediparra. The modern ending for an avian suborder is -ida, so it become Parrida. Unlike the names of family and genus level groups, names ordinal level groups are not regulated by the International Commission on Zoological Names. Nonetheless, I continue to follow the principle of priority and as far as I can tell, Sharpe was the first to name this group.
Parrida consists of 4 small families totalling 16 species. In spite of the small number of species, the Parrida are futher divided into two superfamilies. The Parrida are sister to the much larger group of sandpipers, which are ranked as both a superfamily (Scolopacoidea) and family (Scolopacidae). Their mutual age is constrained above and below by calibration points, and the estimated age of both parvorders is 49.3 million years. The divisions between the small groups contained in Parrida are deep enough for family recognition. Moreover, each family is easily recognizable.
family and genus tree
Černý and Natale's (2021) calibration points are fairly close here, so the ages should be reasonably accurate. Reading off their tree, the division between the Plains-wanderer and the seedsnipes occurred 28 mya, and the division between the painted-snipes and jacanas occurred 34 mya, both in the Oligocene These numbers are a bit older than the calibration points of 30.5 and 24.5 mya, respectively. The Parrida's common ancestor probably lived in the Eocene epoch, about 43 mya.
Thinocoroidea Sundevall, 1836
The Thinocoroidea's origin dates back about 38 million years. It consists of the monotypic plains-wanderer family (Pedionomidae) and the seedsnipe family (Thinocoridae). The two families seem to be separated by about 28.7 million years.
Pedionomidae: Plains-wanderer Bonaparte, 1856
1 genus, 1 species HBW-3
- Plains-wanderer, Pedionomus torquatus
Thinocoridae: Seedsnipes Sundevall, 1836
2 genera, 4 species HBW-3
- Rufous-bellied Seedsnipe, Attagis gayi
- White-bellied Seedsnipe, Attagis malouinus
- Gray-breasted Seedsnipe, Thinocorus orbignyianus
- Least Seedsnipe, Thinocorus rumicivorus
Jacanoidea Chenu & des Murs, 1854 (1840)
Like the Thinocoroidea, the Jacanoidea are about 38 million years old. The two included families, the painted-snipes (Rostratulidae) and jacanas (Jacanidae) separated about 33.7 mya.
Rostratulidae: Painted-snipes Mathews, 1913-14 (1855)
2 genera, 3 species HBW-3
- South American Painted-snipe, Nycticryphes semicollaris
- Greater Painted-snipe, Rostratula benghalensis
- Australian Painted-snipe, Rostratula australis
Jacanidae: Jacanas Chenu & des Murs, 1854 (1840)
6 genera, 8 species HBW-3
|Click for Jacanidae|
Whittingham et al. (2000) found two clades of Jacanas, one containing Hydrophasianus and Jacana, the other consisting of the other four genera.
- Pheasant-tailed Jacana, Hydrophasianus chirurgus
- Northern Jacana, Jacana spinosa
- Wattled Jacana, Jacana jacana
- African Jacana, Actophilornis africanus
- Madagascan Jacana, Actophilornis albinucha
- Bronze-winged Jacana, Metopidius indicus
- Lesser Jacana, Microparra capensis
- Comb-crested Jacana, Irediparra gallinacea
Scolopaci Strauch, 1978
The parorder Scolopaci originated over 49 million years ago. It contains only one superfamily, Scolopacoidea.
Scolopacoidea Rafinesque, 1815
The superfamily Scolopacoidea has the same age as Scolopaci, 49.3 million years. It currently contains one family, which is divided into 5 subfamilies. If this age holds up, we may to promote some of those subfamilies to families.
Scolopacidae: Sandpipers, Snipes Rafinesque, 1815
28 genera, 97 species HBW-3
|Click for Scolopacidae|
The overall treatment of the sandpipers is based on Černý and Natale (2021). It is generally consistent with the results in Baker et al. (2007), Ericson et al. (2003a), Fain and Houde (2007), Gibson (2010), Gibson and Baker (2012), Paton et al. (2003), Pereira and Baker (2005), and Thomas et al. (2004a) as well the more heterogeneous evidence assembled by Thomas et al. (2004b).
Černý and Natale are the first to include DNA from the Slender-billed Curlew and Nordmann's Greenshank.
The curlews and Upland Sandpiper form the basal subfamily (Numeninae) with an estimated age of 42.7 million years. If that holds up, a promotion to family is in order. The next branch is the godwits (Limosinae) at about 40.5 million years---another potential family. After this come the dowitchers, snipe, and woodcock (Scolopacinae), phalaropes and shanks (Tringinae), and finally turnstones and stints (Arenariinae) as in the diagram. The ages are estimated at 37.3, 36, and 36 million years, respectively---again each is old enough for family status.
The dowitchers, snipes, and woodcocks (Scolopacinae) are next.
Lymnocryptes Issues: The sample of the Jack Snipe (Lymnocryptes) used by Černý and Natale (2021) seems to be a chimera, so I have ignored their results for it. Baker et al. (2007) found Lymnocryptes sister to a dowitcher (they used Limnodromus scolopaceus), so I have put it there.
New Zealand Snipe: The New Zealand Snipes (Coenocorypha) are now considered to include 5 exant and recently extinct species based on Baker et al. (2010) and Worthy et al. (2002). The Snares and South Island Snipes are quite closely related, with an estimated divergence time of about 50,000 years. Their status as separate species rests on the lack of an aerial display for the Snares Snipe, as well as genetic and plumage differences. The other Coenocorypha are somewhat more distant relatives.
Gibson (2010) and Gibson and Baker (2012) found that Imperial Snipe is more closely related to the New Zealand snipes than to the other snipes. The Imperial Snipe and two other snipes have sometimes been separated as Chubbia (Mathews 1913), and that is how I treat them here. It is clear that this arrangement of the snipes is not fully satisfactory, and they deserve further study.
Gallinago Split: Černý and Natale (2021) made clear there is a deep division in Gallinago, which I recognize by putting them in separate genera.
- The Great Snipe, Gallinago media, Solitary Snipe, Gallinago solitaria, Wood Snipe, Gallinago nemoricola, Swinhoe's Snipe, Gallinago megala, Pin-tailed Snipe, Gallinago stenura, and Latham's Snipe, Gallinago hardwickii are all transferred to genus Telmatias (Boie 1826, type stenura).
- The other 8 Gallinago snipes remain in Gallinago.
Tribes in Tringidae: The dates in Tringidae are rather unreliable as there are no nearby calibration points. Ignoring that, and that the calibrated Černý and Natale (2021) tree differs a litte from the TiF tree, we have two obvious breakpoints. The phalaropes and Terek Sandpiper form a group, and the Common and Spotted Sandpipers are well-separated from the rest of Tringidae. The tribes are then Phalaropodini (Bonaparte 1831), Actitini (Informal 2021), and Tringini (Rafinesque 1815).
Willets: The Willet, Tringa semipalmata, has been split into Western Willet, Tringa inornata, and Eastern Willet, Tringa semipalmata, based on Oswald et al., (2016).
Tringa Split: Černý and Natale (2021) provide a better-calibrated tree than past efforts. There's not a good calibration point near the Tringinae, so I assume ages are substantially over-estimated. Nonetheless, it provides some guidence concerning where to draw genus boundaries. The previous list had used three subgenera for Tringa and I'm promoting them to regular genera. As a result,
- Tringa (Linnaeus 1758, type ochropus) is restricted to the Green Sandpiper, Tringa ochropus, and the Solitary Sandpiper, Tringa solitaria
- The two tattlers are transferred to genus Heteroscelus (Baird 1858, type brevipes)
- The remaining Tringa sandpipers are placed in genus Totanus (Bechstein 1803, type totanus).
Prosobonia: Cibois et al. (2012) found that Aechmorhynchus and Prosobonia are very closely related, enough so to merge Aechmorhynchus (Coues 1874, type parvirostris) into Prosobonia (Bonaparte 1850, type leucoptera). Černý and Natale (2021) have it as the basal group in Arenariinae, and I have put it in tribe Prosoboniini (Bonaparte, 1850).
Aphriza merged into Calidris: It has long been suspected the Surfbird is close to the knots (e.g., Jehl, 1968). This is exactly what Bororwik and McLennan (1999) found in their DNA tree. Indeed, their results suggest the Surfbird and knots are congeneric. The recent analysis by Gibson (2010) and Gibson and Baker (2012), using additional data, concured, as did Černý and Natale (2021). Based on this, I've merged Aphriza into Calidris.
Tribes in Arenariinae: Černý and Natale (2021) have a calibration point that somewhat constrains dates for the broad Calidris sandpipers. To me, it suggests dividing the subfamily Arenariinae into three tribes: Prosoboniini (Bonaparte 1850), Arenariini, and Calidrini (Reichenbach 1849 (1838)). This creates the handy term Calidrine, which can be used to refer to the species formerly in the very broad Calidris.
Černý and Natale's results alao suggest that the broad genus Calidris should be split into a number of genera. I had previously done such a thing, and then undone it after the AOU, BOU, and H&M 4th ed. have all merged all of these species into Calidris, as in Banks (2012).
Well, I'm undoing it and then some!
- Calidris (Merrem 1804, type canutus) is restricted to three species: Great Knot, Calidris tenuirostris, Red Knot, Calidris canutus, and Surfbird, Calidris virgata.
- The Ruff, Calidris pugnax, is transferred to Philomachus (Merrem 1804, monotypic).
- Broad-billed Sandpiper, Calidris falcinellus, and Sharp-tailed Sandpiper, Calidris acuminatus are transferred to Limicola (Kaup 1816, type falcinellus).
- Curlew Sandpiper, Calidris ferruginea is transferred to the monotypic genus Erolia (Vieillot 1816).
- Stilt Sandpiper, Calidris himantopus is transferred to the monotypic genus Micropalama (Baird 1858).
- Temminck’s Stint, Calidris temminckii, Long-toed Stint, Calidris subminutus, Red-necked Stint, Calidris ruficollis, and Spoon-billed Sandpiper, Calidris pygmeus are placed in Eurynorhynchus (Nilsson 1821, type pygmeus).
- Buff-breasted Sandpiper, Calidris subruficollis is placed in the monotypic genus Tryngites (Cabanis 1856).
- Sanderling, Calidris alba, Dunlin, Calidris alpina, Purple Sandpiper, Calidris maritima, and Rock Sandpiper, Calidris ptilocnemis are transferred to Pelidna (Cuvier 1816, type alpina).
- Baird's Sandpiper, Calidris bairdii, Little Stint, Calidris minuta, Least Sandpiper, Calidris minutilla, White-rumped Sandpiper, Calidris fuscicollis, Pectoral Sandpiper, Calidris melanotos, Semipalmated Sandpiper, Calidris pusilla, and Western Sandpiper, Calidris mauri are transferred to Ereunetes (Illiger 1811, type pusilla).
It's interesting that the Eurasian stints mostly end up in Eurynorhynchus while the similarly-sized American peeps are in Ereunetes. Notice that the Little Stint is in the Amrican peep group.
Numeniinae: Curlews G.R. Gray, 1840
- Upland Sandpiper, Bartramia longicauda
- Bristle-thighed Curlew, Numenius tahitiensis
- Whimbrel, Numenius phaeopus
- Little Curlew, Numenius minutus
- Eskimo Curlew, Numenius borealis
- Slender-billed Curlew, Numenius tenuirostris
- Long-billed Curlew, Numenius americanus
- Far Eastern Curlew, Numenius madagascariensis
- Eurasian Curlew, Numenius arquata
Limosinae: Godwits G.R. Gray, 1841
- Bar-tailed Godwit, Limosa lapponica
- Black-tailed Godwit, Limosa limosa
- Hudsonian Godwit, Limosa haemastica
- Marbled Godwit, Limosa fedoa
Scolopacinae: Dowitchers, Snipe, and Woodcock Rafinesque, 1815
- Jack Snipe, Lymnocryptes minimus
- Asian Dowitcher, Limnodromus semipalmatus
- Short-billed Dowitcher, Limnodromus griseus
- Long-billed Dowitcher, Limnodromus scolopaceus
- Eurasian Woodcock, Scolopax rusticola
- American Woodcock, Scolopax minor
- Amami Woodcock, Scolopax mira
- Javan Woodcock, Scolopax saturata
- New Guinea Woodcock, Scolopax rosenbergii
- Bukidnon Woodcock, Scolopax bukidnonensis
- Sulawesi Woodcock, Scolopax celebensis
- Moluccan Woodcock, Scolopax rochussenii
- Imperial Snipe, Chubbia imperialis
- Jameson's Snipe, Chubbia jamesoni
- Fuegian Snipe, Chubbia stricklandii
- †North Island Snipe, Coenocorypha barrierensis
- Subantarctic Snipe, Coenocorypha aucklandica
- Chatham Snipe, Coenocorypha pusilla
- †South Island Snipe, Coenocorypha iredalei
- Snares Snipe, Coenocorypha huegeli
- Great Snipe, Telmatias medius
- Wood Snipe, Telmatias nemoricola
- Solitary Snipe, Telmatias solitarius
- Swinhoe's Snipe, Telmatias megalus
- Pin-tailed Snipe, Telmatias stenurus
- Latham's Snipe, Telmatias hardwickii
- Giant Snipe, Gallinago undulata
- Noble Snipe, Gallinago nobilis
- South American Snipe, Gallinago paraguaiae
- Puna Snipe, Gallinago andina
- African Snipe, Gallinago nigripennis
- Madagascan Snipe, Gallinago macrodactyla
- Common Snipe, Gallinago gallinago
- Wilson's Snipe, Gallinago delicata
Tringinae: Phalaropes and Shanks Rafinesque, 1815
- Terek Sandpiper, Xenus cinereus
- Wilson's Phalarope, Steganopus tricolor
- Red-necked Phalarope, Phalaropus lobatus
- Red Phalarope / Gray Phalarope, Phalaropus fulicarius
- Common Sandpiper, Actitis hypoleucos
- Spotted Sandpiper, Actitis macularius
- Green Sandpiper, Tringa ochropus
- Solitary Sandpiper, Tringa solitaria
- Gray-tailed Tattler, Heteroscelus brevipes
- Wandering Tattler, Heteroscelus incanus
- Marsh Sandpiper, Totanus stagnatilis
- Wood Sandpiper, Totanus glareola
- Common Redshank, Totanus totanus
- Lesser Yellowlegs, Totanus flavipes
- Spotted Redshank, Totanus erythropus
- Greater Yellowlegs, Totanus melanoleuca
- Common Greenshank, Totanus nebularia
- Nordmann's Greenshank, Totanus guttifer
- Western Willet, Totanus inornata
- Eastern Willet, Totanus semipalmata
Arenariinae: Turnstones and Stints Stejneger, 1885 (1840)
- †Kiritimati Sandpiper, Prosobonia cancellata
- Tuamotu Sandpiper, Prosobonia parvirostris
- †Tahiti Sandpiper, Prosobonia leucoptera
- †Moorea Sandpiper, Prosobonia ellisi
- Ruddy Turnstone, Arenaria interpres
- Black Turnstone, Arenaria melanocephala
- Great Knot, Calidris tenuirostris
- Red Knot, Calidris canutus
- Surfbird, Calidris virgata
- Ruff, Philomachus pugnax
- Broad-billed Sandpiper, Limicola falcinellus
- Sharp-tailed Sandpiper, Limicola acuminata
- Curlew Sandpiper, Erolia ferruginea
- Stilt Sandpiper, Micropalama himantopus
- Temminck's Stint, Eurynorhynchus temminckii
- Long-toed Stint, Eurynorhynchus subminuta
- Red-necked Stint, Eurynorhynchus ruficollis
- Spoon-billed Sandpiper, Eurynorhynchus pygmeus
- Buff-breasted Sandpiper, Tryngites subruficollis
- Sanderling, Pelidna alba
- Dunlin, Pelidna alpina
- Purple Sandpiper, Pelidna maritima
- Rock Sandpiper, Pelidna ptilocnemis
- Baird's Sandpiper, Ereunetes bairdii
- Little Stint, Ereunetes minutus
- Least Sandpiper, Ereunetes minutillus
- White-rumped Sandpiper, Ereunetes fuscicollis
- Pectoral Sandpiper, Ereunetes melanotos
- Semipalmated Sandpiper, Ereunetes pusillus
- Western Sandpiper, Ereunetes mauri
Turnici Huxley, 1868
The buttonquail (Turnici) are sister to the Lari. For a long time, they weren't considered to be related to shorebirds. They were often grouped with the plains-wanderer and put in the Gruiformes. Although the Monroe and Sibley (1993) checklist removed them from the Gruiformes, they were placed in their own parvclass, with no close relative.
In their pioneering study of the Charadriiformes, Paton et al. (2003) attempted to use Turnix as an outgroup. After all, van Tuinen et al. (2000) had Turnix outside the shorebirds, but close enough to make a useful outgroup. They got a surprise! They found the buttonquail were well-embedded in the Charadriiformes, sister to the group I now call Lari. And as other studies have also found, the division between the Turnici and Lari appeared quite deep. That's why I maintain them as distinct suborders.
Turnicidae: Buttonquail G.R. Gray, 1840 (1831)
2 genera, 17 species HBW-3
|Click for Turnicidae|
The buttonquail tree was constructed by combining the species groups described by Debus (1996) with the 7-species phylogeny of Černý and Natale (2021). I was surprised there were no conflicts. The fact that there weren't reminds us that traditional methods often did a pretty good job of determining the phylogeny. Debus also describes the morphological differences between quail and buttonquail that justify grouping them together, including Ortyxelos. See also the discussion in Sibley and Ahlquist (1990).
I do have a reservation about the resulting phylogeny. It seems rather odd to have two African species (Hottentot and Black-rumped) embedded in an Australasian clade.
- Quail-plover, Ortyxelos meiffrenii
- Yellow-legged Buttonquail, Turnix tanki
- Spotted Buttonquail, Turnix ocellatus
- Barred Buttonquail, Turnix suscitator
- Madagascan Buttonquail, Turnix nigricollis
- Common Buttonquail, Turnix sylvaticus
- Red-backed Buttonquail, Turnix maculosus
- Little Buttonquail, Turnix velox
- Worcester's Buttonquail, Turnix worcesteri
- Sumba Buttonquail, Turnix everetti
- Red-chested Buttonquail, Turnix pyrrhothorax
- Hottentot Buttonquail, Turnix hottentottus
- Black-rumped Buttonquail, Turnix nanus
- Black-breasted Buttonquail, Turnix melanogaster
- Painted Buttonquail, Turnix varius
- Chestnut-backed Buttonquail, Turnix castanotus
- Buff-breasted Buttonquail, Turnix olivii
Lari Sharpe 1891
The Lari include the gulls and terns, but also more. They contain two parvorders, Glareolida (crab plover, coursers, pratincoles), and Larida (skuas, auks, gulls, terns, skimmers). The two groups seem to have split around 44 mya, hence the parvorder status, but there is some uncertainty about the timing as much of the Lari tree lacks calibration points.
Glareolida Sharpe, 1891
The Crab Plover's nearest relatives are the coursers and pratincoles, the Glareolidae. There's a fairly small time gap between the two families in Glareolida on Černý and Natale (2021) tree, about 2.7 million years, with ages of 40 million years or so. But there's no relevant calibration point on this section of the tree, so the ages are likely quite exaggerated. While the gap is small, the time depth is sufficient for family status for both. So I keep two separate families here.
Dromadidae: Crab Plover G.R. Gray, 1840
1 genus, 1 species HBW-3
Pereira and Baker (2010) found that Dromas is sister to the Glareolidae, as represented by Glareola and Cursorius.
- Crab Plover, Dromas ardeola
Glareolidae: Coursers, Pratincoles Brehm, 1831
3 genera, 17 species HBW-3
|Click for Courser and|
The arrangement is based on the genetic analyses of Cohen (2011), which uses up to four genes (ND2, Fib5, TGFB, GAPDH), and of Černý and Natale (2021). The result is as consistent as I can make it with Figures A.5 (RAxML) and A.7 (ExaBayes) in Černý and Natale and with Figure 4.3 in Cohen (nuclear + mitochondrial DNA). I've appealed to the non-genetic analyses in those papers, together with what HBW says about their relations, and just plain looking at pictures and range maps.
As a result, we now have 7 genera instead of 3. Since there are no calibration anywhere near Glareolidae, and since the total evidence tree in Černý and Natale (2021) is so different from what I am using, it doesn't provide much help. I did make some guesses about time intervals based on the DNA trees, but mainly focused on making each genus a coherent group. Here are the generic changes:
- The Double-banded Courser, previously Rhinoptilus africanus, has been transferred to Smutsornis (Roberts 1922, monotypic).
- The Gray Pratincole, Glareola cinerea, and Small Pratincole, Glareola lactea have been transferred to Galachrysia (Bonaparte 1856, type lactea). The movement that prompted this was based on DNA analyzed by Černý and Natale (2021).
- The Rock Pratincole, Glareola nuchalis, and Madagascan Pratincole, Glareola ocularis, have been transferred to Subglareola (Mathews 1913, type ocularis).
- I have restored the monotypic genus Stiltia (G.R. Gray, 1855, type isabella) which I had previously submerged in Glareola.
Glareolidae: Coursers, Pratincoles
- Three-banded Courser, Rhinoptilus cinctus
- Bronze-winged Courser, Rhinoptilus chalcopterus
- Jerdon's Courser, Rhinoptilus bitorquatus
- Double-banded Courser, Smutsornis africanus
- Gray Pratincole, Galachrysia cinerea
- Small Pratincole, Galachrysia lactea
- Cream-colored Courser, Cursorius cursor
- Somali Courser, Cursorius somalensis
- Burchell's Courser, Cursorius rufus
- Temminck's Courser, Cursorius temminckii
- Indian Courser, Cursorius coromandelicus
- Rock Pratincole, Subglareola nuchalis
- Madagascan Pratincole, Subglareola ocularis
- Australian Pratincole, Stiltia isabella
- Black-winged Pratincole, Glareola nordmanni
- Collared Pratincole, Glareola pratincola
- Oriental Pratincole, Glareola maldivarum
larida Rafinesque, 1815
The Larida contain two superfamilies, the Alcoidea (skuas and auks) and the Laroidea (skimmers, terns, gulls).
Alcoidea Leach, 1820
Stercorariidae: Skuas, Jaegers G.R. Gray, 1870 (1831)
1 genus, 7 species HBW-3
The skuas were formerly separated in the genus Catharacta, but a genetic study by Cohen et al. (1997) found the Great Skua and Pomarine Jaeger were sister species. This has been called into question by Braun and Brumfield (1998), who argued that all the skuas might be sister to the Pomarine Jaeger. Either way, the 2-genus treatment was untenable, and most have decided to merge Catharacta into Stercorarius. If Braun and Brumfield are right, an alternative would be to put the Pomarine Jaeger by itself in Coprotheres and the skuas in Catharacta. This doesn't work in the current treatment, which is slightly different, and is based on Černý and Natale (2021).
The species boundaries in the southern skuas are somewhat uncertain. See Ritz et al. (2008).
- Long-tailed Jaeger, Stercorarius longicaudus
- Parasitic Jaeger / Arctic Skua, Stercorarius parasiticus
- Pomarine Jaeger / Pomarine Skua, Stercorarius pomarinus
- Great Skua, Stercorarius skua
- Brown Skua, Stercorarius antarcticus
- Chilean Skua, Stercorarius chilensis
- South Polar Skua, Stercorarius maccormicki
Alcidae: Auks Leach, 1820
12 genera, 25 species HBW-3
|Click for Alcidae|
The taxonomy follows Baker et al. (2007) and Pereira and Baker (2008). The time scale in Pereira and Baker should be taken with a grain of salt. Considering that some of the fossils used as calibration points are distantly related, or of controversial affinities, perhaps with a whole trunkload of salt would be better.
Humphries and Winker (2010) clarified some details about Aethia.
Xantus's Murrelet Split: For the split of Scripps's Murrelet, Synthliboramphus scrippsi, from Xantus's Murrelet, Synthliboramphus hypoleucus, see Birt et al. (2012). Further, Synthliboramphus hypoleucus takes the name Guadalupe Murrelet.
Tropical Murrelets: Finally, based on the dates from Černý and Natale (2021), I've moved the tropical murrelets to genus Endomychura (Oberholser 1899, type hypoleuca).
Aethiinae AOU 1908 (1840)
- Rhinoceros Auklet, Cerorhinca monocerata
- Tufted Puffin, Fratercula cirrhata
- Atlantic Puffin, Fratercula arctica
- Horned Puffin, Fratercula corniculata
- Cassin's Auklet, Ptychoramphus aleuticus
- Crested Auklet, Aethia cristatella
- Least Auklet, Aethia pusilla
- Whiskered Auklet, Aethia pygmaea
- Parakeet Auklet, Aethia psittacula
Alcinae Leach 1820
- Long-billed Murrelet, Brachyramphus perdix
- Marbled Murrelet, Brachyramphus marmoratus
- Kittlitz's Murrelet, Brachyramphus brevirostris
- Black Guillemot, Cepphus grylle
- Spectacled Guillemot, Cepphus carbo
- Pigeon Guillemot, Cepphus columba
- Thick-billed Murre / Brunnich's Guillemot, Uria lomvia
- Common Murre / Guillemot, Uria aalge
- Dovekie / Little Auk, Alle alle
- Razorbill, Alca torda
- †Great Auk, Pinguinus impennis
- Ancient Murrelet, Synthliboramphus antiquus
- Japanese Murrelet, Synthliboramphus wumizusume
- Craveri's Murrelet, Endomychura craveri
- Scripps's Murrelet, Endomychura scrippsi
- Guadalupe Murrelet, Endomychura hypoleuca
Laroidea Rafinesque, 1815
The exact placement of several genera (Rynchops, Anous, Gygis) remains somewhat problematic. It's clear that they are old basal lineages within Laroidea, but exactly where and how they fit with Sternidae and Laridae remains uncertain. The current arrangement derives from Černý and Natale (2021), placing Rhynchops and Gygis in their own basal subfamilies in Sternidae, and Anous as the basal subfamily in Laroidea. If you only looked at the main paper, you won't understand how I can say that. The answer is in Appendix A, which contains the purely genetic trees. Both the ASTRAL III and RAxML trees (Figures A-3/4 and A-5/6, respectively) agree with this placement of Anous. However, the ExaBayes tree (Figs. A-7/8) disagrees.
There are four older relevant papers: Bridge et al. (2005), Baker et al. (2007), Ödeen et al. (2010), and Jackson et al. (2012). They successively analyzed more genes, but Baker et al. has better taxon sampling in the terns. From those papers, it was already clear that the skimmers are by themselves, the gulls group together, and the terns other than Anous and Gygis also form a clade. Whether all of the terns are monophyletic, or whether the core terns plus Gygis are monophyletic, was not resolved, and remains so today.
Bridge et al. (2005) placed Anous and Gygis in the Sternidae. In contrast, Baker et al. (2007) found Anous and Gygis to be basal to the gulls, terns, and skimmers. In their combined analysis, Ödeen et al (2010) group Anous and Gygis basally in Laridae, with a trichotomy between the skimmers, gulls, and remaining terns. They note that this arrangement is weakly supported. They also give a somewhat better supported mitochondrial tree which puts Gygis with the other terns and Anous with the gulls and skimmers. Finally, Jackson et al. (2012) consider more genes, but leave out the noddies. The resulting combined analysis puts the white terns sister to the terns, and the skimmers as basal group. The mitochondrial DNA results are a bit different, and put the white terns as basal, then skimmers, then gulls and terns.
This suggests that we think of the larids as consisting of five main groups: gulls, terns, skimmers, noddies, and white terns. I emphasize this by ranking each group as a subfamily. There's no consensus about how they fit together. I was treating them as a five-way polytomy. However, I'm now using the results from Figs A-3 to A-6 from Černý and Natale (2021). This is the most comprehensive analysis. It groups the white terns and skimmers with the terns, and the noddies with the gulls.
Sternidae: Terns and Skimmers Vigors, 182s5
11 genera, 44 species HBW-3
|Click for Sternidae|
Rynchopinae: Skimmers Bonaparte, 1838
- African Skimmer, Rynchops flavirostris
- Indian Skimmer, Rynchops albicollis
- Black Skimmer, Rynchops niger
Gyginae: White Terns Verheyen, 1959
- White Tern, Gygis alba
- Little White-Tern, Gygis microrhyncha
Sterninae: Terns Vigors, 1825
There are differences in the tern taxonomy in Bridge et al. (2005) and Baker et al. (2007). I've adopted the Bridge et al. framework because they sampled many more tern species than Baker et al. Use of Černý and Natale (2021) hasn't changed much here.
Cabot's Tern: The American Sandwich Terns are split under the old name Cabot's Tern, Thalasseus acuflavidus. Efe et al. (2009) found that Cabot's Tern is more closely related to Elegant Tern than to Old World Sandwich Terns. They also found that no systematic genetic distinction between Cayenne and Cabot's Terns and question whether eurygnathus is distinct from acuflavidus.
Sterninae: Terns Vigors, 1825
- Aleutian Tern, Onychoprion aleuticus
- Sooty Tern, Onychoprion fuscatus
- Gray-backed Tern / Spectacled Tern, Onychoprion lunatus
- Bridled Tern, Onychoprion anaethetus
- Little Tern, Sternula albifrons
- Least Tern, Sternula antillarum
- Fairy Tern, Sternula nereis
- Yellow-billed Tern, Sternula superciliaris
- Saunders's Tern, Sternula saundersi
- Damara Tern, Sternula balaenarum
- Peruvian Tern, Sternula lorata
- Large-billed Tern, Phaetusa simplex
- Caspian Tern, Hydroprogne caspia
- Gull-billed Tern, Gelochelidon nilotica
- Inca Tern, Larosterna inca
- Whiskered Tern, Chlidonias hybrida
- Black-fronted Tern, Chlidonias albostriatus
- White-winged Tern, Chlidonias leucopterus
- Black Tern, Chlidonias niger
- Sandwich Tern, Thalasseus sandvicensis
- Elegant Tern, Thalasseus elegans
- Cabot's Tern, Thalasseus acuflavidus
- Great Crested Tern / Greater Crested Tern, Thalasseus bergii
- Chinese Crested Tern, Thalasseus bernsteini
- Lesser Crested Tern, Thalasseus bengalensis
- Royal Tern, Thalasseus maximus
- Forster's Tern, Sterna forsteri
- Snowy-crowned Tern, Sterna trudeaui
- Arctic Tern, Sterna paradisaea
- South American Tern, Sterna hirundinacea
- Kerguelen Tern, Sterna virgata
- Antarctic Tern, Sterna vittata
- Common Tern, Sterna hirundo
- Black-naped Tern, Sterna sumatrana
- Roseate Tern, Sterna dougallii
- White-fronted Tern, Sterna striata
- White-cheeked Tern, Sterna repressa
- River Tern, Sterna aurantia
- Black-bellied Tern, Sterna acuticauda
Laridae: Gulls and Noddies Rafinesque, 1815
15 genera, 62 species HBW-3
|Click for Laridae|
Anoinae: Noddies Bonaparte, 1854
As mentioned above, two of the three genetic analyses in Černý and Natale (2021, Figs. A-3/4, A-5) found the noddies sister to the gulls. We rank them as the subfamily Anoinae. The Noddy portion of the tree is from Cibois et al. (2016), who found that Procelsterna is embedded in Anous. Since the noddy clade is of recent origin, it makes sense to treat them all as a single genus. Anous (Stephens 1826, type stolidus) has priority over Procelsterna (Lafresnaye 1842, type ceruleus teretriostris).
- Brown Noddy, Anous stolidus
- Black Noddy, Anous minutus
- Lesser Noddy, Anous tenuirostris
- Blue-gray Noddy / Blue Noddy, Anous ceruleus
- Gray Noddy, Anous albivitta
Larinae: Gulls Rafinesque, 1815
At the generic level, the present TiF taxonomy of the gulls is a refinement of Sternkopf (2011) and Pons et al. (2005). Sternkopf used a wider variety of genes than Pons et al., but cut back a bit on taxa considered. Their results are generally consistent with each other and with Crochet et al., (2000). The Pons et al. treatment was quickly accepted by AOU and BOU. The recent paper by Černý and Natale (2021) increased certainty about some of the nodes, and broke ties in a couple of cases.
The use of Sternkopf (2011) and Černý and Natale (2021) means that the Little/Ross's Gull clade is one of the basal groups and that the Ring-billed and Mew Gulls are sister species. It's also led to some minor reordering within genera. Although Sternkopf did not consider Saunder's Gull, Černý and Natale (2021) did, its current position on a branch just before Chroicocephalus reflects that.
As recently as H&M-3, most gulls were put in genus Larus. The exceptions were pretty clearly different: Swallow-tailed Gull (Creagrus), Ross's Gull (Rhodostethia), Ivory Gull (Pagophila), Sabine's Gull (Xema), the two kittiwakes (Rissa), and the Dolphin Gull (Leucophaeus). Everything else was in Larus.
This classification started to change once the DNA was studied. Pons et al. (2005) started the changes with by noting the various types of gulls on their consensus tree (Fig. 2). Besides those listed above (except Dolphin Gull), they separated Saunder's Gull, the masked gulls, hooded gulls, black-headed gulls, band-tailed gulls, and white-headed gulls. Pretty soon, Saunder's Gull moved to genus Saundersilarus, the masked gulls to Chroicocephalus, hooded gulls and Dolphin Gull to Leucophaeus, black-headed to Ichthyaetus, and band-tailed and white-headed gulls remained in Larus, as in H&M-4.
New Genera: Consideration of Černý and Natale (2021) allows some refinement of this arrangement.
- The Slender-billed Gull, Chroicocephalus genei, is separated from the other Chroicocephalus by about 10 million years. It is also distinctive, and I've moved it to genus Gelastes (Bonaparte 1856, monotypic).
- The division between Dolphin and Gray Gulls on one hand, and Laughing, Franklin's, and Lava Gulls on the other is about 6 million years, enough to support different genera. Moreover, they obviously form two, or even three groups. Accordingly, the Laughing, Franklin's and Lava Gulls have been moved to Atricilla (Bonaparte 1854, type atricilla). It would not be unreasonable to also split the Gray Gull, Leucophaeus modestus. In that case it would be genus Blasipus (Bruch 1853).
- The band-tailed gulls (4 species) are transferred to genus Gabianus (Bruch 1853, type pacificus).
When it comes to Larus, we have more information, and we need it! The papers by Liebers et al. (2001, 2002, 2004), de Knijff et al. (2001), Crochet et al. (2002), Pons et al. (2004), and Gay et al. (2005) focused on the big white-headed gulls. They also get full coverage by Sternkopf (2011) and Černý and Natale (2021).
In spite of all the above effort, Larus gull relationships remain somewhat obscured by past and recent hybridization. Questions about possible ring species have been raised, and denied (Liebers et al., 2004). Species limits within the herring gulls remain controversial. Nonetheless, a coherent picture is gradually being teased out.
The TiF list has adopted Figs. A-5 and A-6 from Černý and Natale for the Larus gulls.
It now seems that Heerman's Gull, Larus heermanni is the first branch of the Larus tree. You'll notice it has a full black tail, different from both the band-tailed gulls — the nearest relatives of Larus — and the remaining Larus gulls. The second branch is a small clade consisting of smaller white-headed gulls: Ring-billed (delawarensis), Common (canus), and Short-tailed (brachyrhynchus) Gulls. Then we get to the big white-headed gulls.
There's considerable support for the Yellow-footed Gull, Larus livens, and Western Gull, Larus occidentalis (both races) being likely sisters, and being the sister group to the main group of Larus gulls. Some of the Larus frequently hybridize, even when far apart on the tree. A prime example is the Western Gull, Larus occidentalis, which hybridizes freely with the Glaucous-winged Gull, Larus glaucescens
The Kelp Gull, Larus dominicanus, is next, in a group by itself. The remaining Larus gulls form two groups, one contains the European Herring Gull, Larus argentatus, the Lesser Black-backed Gull, Larus fuscus, and some close relatives. The other includes the Great Black-backed Gull, Larus marinus, the American Herring Gull, Larus smithsonianus, Iceland Gull, Larus glaucoides, and the far eastern herring-type gulls.
Laridae — Further Taxonomic Notes
Chroicocephalus: I'm now primarily using Fig. A-7 of Černý and Natale (2021) for Chroicocephalus with a little bit of Given et al. (2005) for the Gray-headed Gull, Chroicocephalus poiocephalus. Given et al.'s arrangement for Chroicocephalus is somewhat different from Černý and Natale's. I'm uncertain which phylogeny for Chroicocephalus is more accurate.
Gray-hooded/headed Gull: The Gray-hooded Gull / Gray-headed Gull, Chroicocephalus cirrocephalus, has been split into the Gray-hooded Gull, Chroicocephalus cirrocephalus, of South America and the Gray-headed Gull, Chroicocephalus poiocephalus, of Africa. Given et al. (2005) found that C. poiocephalus was more closely related to Hartlaub's Gull, Chroicocephalus hartlaubii than to C. cirrocephalus. They noted this relationship could be an artifact of introgression, but they did not have any evidence to that effect.
White-eyed Gull: I gave the White-eyed Gull, Ichthyaetus leucophthalmus, a question mark even though its DNA was sampled by Pons et al. (2005). The problem is that the arrangement of Ichthyaetus in Pons et al. is quite different from the one used here, and I cannot properly interpret the results. There's no doubt its in the black-headed gull clade, but where it goes is rather uncertain. I thought putting it sister to the Sooty Gull, Ichthyaetus hemprichii made the most sense. Be warned that this choice does not have a lot of data behind it.
Thayer's Gull Lump: Some of the species limits among the herring gulls remain contentious. E.g., see Pittaway (1999) and Weir et al. (2000) concerning whether glaucoides, kumlieni, and thayeri are one, two, or even three species. For the present, I am following AOU on this (one). So Thayer's Gull, Larus thayeri, is lumped into Iceland Gull, Larus glaucoides.
Possible Splits: Other named taxa in Liebers's clade II sometimes considered species include (West) Siberian Gull (heuglini and maybe taimyrensis), Baltic Gull (fuscus), Steppe Gull (barabensis). They also suggest there are two types of Glaucous Gull which may not match traditional subspecies, and two types of European Herring Gulls that definitely don't match traditional subspecies. There's also a question concerning whether the Yellow-legged Gull should be split into Mediterranean Yellow-legged Gull (michahellis), Atlantic Yellow-legged Gull (atlantis) and possibly even Cantabrican Yellow-legged Gull (lusitanius?).
- Swallow-tailed Gull, Creagrus furcatus
- Little Gull, Hydrocoloeus minutus
- Ross's Gull, Rhodostethia rosea
- Ivory Gull, Pagophila eburnea
- Sabine's Gull, Xema sabini
- Black-legged Kittiwake, Rissa tridactyla
- Red-legged Kittiwake, Rissa brevirostris
- Saunders's Gull, Saundersilarus saundersi
- Slender-billed Gull, Gelastes genei
- Bonaparte's Gull, Chroicocephalus philadelphia
- Black-headed Gull, Chroicocephalus ridibundus
- Brown-headed Gull, Chroicocephalus brunnicephalus
- Gray-hooded Gull, Chroicocephalus cirrocephalus
- Gray-headed Gull, Chroicocephalus poiocephalus
- Hartlaub's Gull, Chroicocephalus hartlaubii
- Red-billed Gull, Chroicocephalus scopulinus
- Silver Gull, Chroicocephalus novaehollandiae
- Black-billed Gull, Chroicocephalus bulleri
- Andean Gull, Chroicocephalus serranus
- Brown-hooded Gull, Chroicocephalus maculipennis
- Gray Gull, Leucophaeus modestus
- Dolphin Gull, Leucophaeus scoresbii
- Laughing Gull, Atricilla atricilla
- Franklin's Gull, Atricilla pipixcan
- Lava Gull, Atricilla fuliginosa
- Pallas's Gull, Ichthyaetus ichthyaetus
- Relict Gull, Ichthyaetus relictus
- Mediterranean Gull, Ichthyaetus melanocephalus
- Audouin's Gull, Ichthyaetus audouinii
- White-eyed Gull, Ichthyaetus leucophthalmus
- Sooty Gull, Ichthyaetus hemprichii
- Pacific Gull, Gabianus pacificus
- Belcher's Gull, Gabianus belcheri
- Black-tailed Gull, Gabianus crassirostris
- Olrog's Gull, Gabianus atlanticus
- Heermann's Gull, Larus heermanni
- Ring-billed Gull, Larus delawarensis
- Common Gull, Larus canus
- Short-billed Gull, Larus brachyrhynchus
- Western Gull, Larus occidentalis
- Yellow-footed Gull, Larus livens
- Kelp Gull, Larus dominicanus
- Glaucous Gull, Larus hyperboreus
- European Herring Gull, Larus argentatus
- Caspian Gull, Larus cachinnans
- Lesser Black-backed Gull, Larus fuscus
- Heuglin's Gull, Larus heuglini
- Great Black-backed Gull, Larus marinus
- Yellow-legged Gull, Larus michahellis
- Armenian Gull, Larus armenicus
- California Gull, Larus californicus
- American Herring Gull, Larus smithsonianus
- Glaucous-winged Gull, Larus glaucescens
- Iceland Gull, Larus glaucoides
- Mongolian Gull, Larus mongolicus
- Vega Gull, Larus vegae
- Slaty-backed Gull, Larus schistisagus