Sylvioidea

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 46 Orders

Paleognathae

Galloanserae

Columbimorphae

Otidimorphae

Strisores

Opisthocomiformes

Gruiformes

Mirandornithes

Ardeae

Charadriiformes

Telluraves

Afroaves

Australaves

Babblers and allies

Babbler overview The babblers have long been treated as a wastebin taxon, with many disparate species dumped in the group. They've sometimes been combined with the Sylviidae, forming an even more disparate group. The previous pages have covered many of the old world warbler families, leaving us with a small sylviid core together with the babblers. The tree to the right is based on Moyle et al. (2012), Fregin et al. (2012, Fig. 1 only), and Gelang et al. (2009). I interpret these papers as evidence in favor of grouping the remaining 450 babblers and allies into 6 families.

One of these families is the new Sylviidae, which has lost most of the old world warblers to new families such as Phylloscopidae and Acrocephalidae. It is sister to the parrotbills, Paradoxornithidae (once known as Panuridae, but Panurus is no longer in this family). The white-eyes remain in an expanded Zosteropidae. Then comes a more narrowly circumscribed Timaliidae, followed by the Pellorneidae. It ends with the laughingthrush family, Leiothrichidae. It would not be unreasonable to rank this last two as subfamilies of Timaliidae, as suggested by Moyle et al. (2012).

The natural name for Leiothrichidae is Turdoididae, which was sometimes used in the early 20th century. However, the ICZN requires keeping separate books for family-level names and for genera, meaning that Leiothrichidae is correct (as are various other family names in current use). I do not know why the ICZN choose this complex system instead of simply extending the genus name system where the oldest-named generic type determines the name. It would be much simpler to also use the oldest-named genus to determine the family name.

Sylviidae: Sylviid Warblers Leach, 1820

3 genera, 37 species HBW-11

In a sense, the Sylviidae are the epicenter of the DNA-driven taxonomic earthquake that has shaken up all of Passeriformes since the 1980's. At times, Sylviidae was considered a huge family, encompassing hundreds of Old World warblers. HBW-11 (del Hoyo et al., 2006), which uses a taxonomy current when the series started in 1992, lists 271 species in Sylviidae. In the TiF list, most of those have been spread across Acrocephalidae, Bernieridae, Cettiidae, Locustellidae, Macrosphenidae, and Phylloscopidae, with a few scattered elsewhere. And that was after HBW had already split off 145 species in Cisticolidae! At times, the babbler family Timaliidae, has also been considered a large family. In HBW-12 (del Hoyo et al., 2007) it contains 309 species.

That was then, this is now. As more genetic studies have been done, it has become clear that Sylviidae's true affinities were with the babblers rather than the other Old World warblers. Cibois (2003) even went so far as to suggest suppressing the name Sylviidae in favor of Timaliidae! Further analysis has clarified matters, and Sylviiade survives, but just barely. It has become a small family.

The changes in Sylvia itself mirror those of the Sylviidae. Most of the Sylvia warblers have been moved to Curruca, leaving only the Blackcap and Garden Warbler. In their place, Sylvia has gained the former Horizorhinus (sometimes considered an Old World flycatcher), and the former Lioptilus and Pseudoalcippe babblers from Timaliidae.

Click for Sylviidae tree
Click for Sylviidae tree

The order here relies on Cibois (2003), and Gelang et al. (2009). In the former two, the Sylviidae are clade 2.

Böhning-Gaese et al. (2003) presented evidence that the Blackcap and Garden Warbler are sister species, close to Pseudoalcippe. Johansson et al. (2008b) found that the Bush Blackcap (Lioptilus) is closer to Pseudoalcippe than to Sylvia. Voelker et al. (2009) found a slightly different topology with Horizorhinus close to Pseudoalcippe. Although Voelker and Light (2011) did not include the Bush Blackcap, they comment on some unpublished work concerning it. Finally, Moyle et al. (2012) included all 4 genera and resovled the issue.

Accordingly, I've submerged Horizonrhinus, Lioptilus, and Pseudoalcippe into Sylvia. The Abyssinian Catbird, Parophasma galinieri, belongs in the clade containing Sylvia, but whether is it belongs in Sylvia or sister to it is uncertain (Gelang et al., 2009). I'm treating it as a sister genus.

Since the type of Sylvia is atricapilla, the remaining Sylvia need a different name. Accordingly I revive the name Curruca for them. Note that the genus Parisoma has been merged into Curruca. The order within Curruca is based on Voelker and Light (2011), which refines previous work by Shirihai et al. (2001), Böhning-Gaese et al. (2003), and Böhning-Gaese et al. (2006).

The arrangement within Curruca is based on Voelker and Light (2011) and, for the curruca complex, Olsson et al. (2013). There are several notable clades which are similar to those described by Shirihai et al. (2001). Voelker and Light estimate the age of the various clades. It would not be unreasonable to divide Curruca into 5 genera. I currently ranked these as subgenera and illustrate them on the tree diagram. The five groups are Atraphornis, Adophoneus, Parisoma, Curruca, and Melizophilus.

The subalpine warbler complex has been split into three (see Brambilla et al., 2008; Svensson 2013). They are Moltoni's Warbler, Curruca subalpina, (also known as moltonii), Western Subalpine Warbler, Curruca inornata, and Eastern Subalpine Warbler, Curruca cantillans. The name subalpina instead of moltonii seems to be preferred for Moltoni's Warbler, although the applicability of subalpina is not clear-cut. See Baccetti et al., (2007) and Svensson (2013).

Olsson et al. (2013) found that except for the already split minula, the “curruca” subspecies east of the Urals (and maybe some birds west of the Urals) are more closely related to Hume's Whitethroat, C. althaea than to curruca. There appear to be at least 4 species here. Although I'd prefer to have claification on things such as range boundaries, songs, and hybridization, I don't. This means that we have 2 choices, either group all 4 clades together as Hume's, or give treat them as species without knowning the whole story. I had originally done the first, but now think the second approach is better as it highlights the problem.

The lesser whitethroat complex includes:

  1. Lesser Whitethroat, Curruca curruca (inc. caucasica)
  2. Desert Whitethroat, Curruca minula
  3. *Siberian Whitethroat, Curruca blythi
  4. *Steppe Whitethroat, Curruca halimodendri (inc. jaxartica)
  5. Mountain Whitethroat / Hume's Whitethroat, Curruca althaea (inc. monticola)
  6. *Stolzmann's Whitethroat, Curruca margelanica (inc. telengitica, chuancheica)

where the asterisks indicate the new species. The English names posed something of a problem. Siberian for blythi was easy enough. The appellations Steppe and Central Asian have been used for halimodendri while althaea goes by Hume's and Mountain, with Mountain being more descriptive. The biggest problem is margelanica. The term Margelanic has been used, based on a city (Margilan) in eastern Uzbekistan. Unfortunately, it is doubly misleading. Margilan is not in the breeding range and Margelanic sounds like it belongs in S. America. Other suggestions include Mongolian or Gobi, but they don't seem quite right either. I thought about Chinese, but that equally applies to minula. The subspecies margelanica was described by Stolzmann, so lacking a better term, I've put his name on it.

Paradoxornithidae: Parrotbills, Fulvettas Horsfield & Moore, 1854

15 genera, 38 species HBW-12

Click for Sylviidae tree
Click for Paradoxornithidae tree

The parrotbills form a small family that are sister to the sylviid warblers (Sylviidae). With one exception, the Wrentit of western North America, they are native to Asia. Since the Sylviidae are primarily birds of Africa and the Palearctic, there is only limited geographic overlap between the two families.

The parrotbill family was previously called Panuridae. However, the genus Panurus (Bearded Reedling, aka Bearded Tit, Bearded Parrotbill) was found to be related to the larks, not to the other parrotbills. The name Paradoxornithidae is next in the priority line, and is applied here.

Unlike its sister clade Sylviidae, Paradoxornithidae has grown. It lost the reedling, but has added the Fulvetta fulvettas (formerly part of Alcippe, see Pasquet et al, 2006; Collar and Robson, 2007), another fulvetta (now Lioparus, formerly Alcippe), a former cisticolid (Rhopophilus), and five babblers: the two Chrysomma, the Wrentit (Chamaea), Moupinia and the highly distinctive Myzornis (Cibois, 2003; Gelang et al. 2009; Moyle et al., 2012).

Between Moyle et al. (2012), Pasquet et al. (2006), and Yeung et al. (2011), we now have a pretty good handle on the phylogeny. Yeung et al. left some uncertainty about whether the Fulvettas belonged in the parrotbill clade, but both Gelang et al. (2009) and Moyle et al. (2012) support the arrangement used here with Fulvetta outside the parrotbill clade.

Yeung et al. (2011) separately considered both nuclear and mitochondrial genes and found four main groups: the Fulvetta group, Neosuthora-Suthora, Chleuasicus-Sinosuthora, and the Paradoxornis-Psittiparus group. However, the nuclear genes studied by Yeung et al. (2011) gave a different arrangement than the mitochondrial genes. The version here groups the small parrotbills together, and treats the fulvetta group as sister to the parrotbills.

Yeung et al.'s (2011) analysis supports splitting the parrotbills into six genera as recommended by Penhallurick and Robson (2009) and Penhallurick (2010). The arrangement of these genera, together with Conostoma and the Wrentit, Chamaea fasciata, follows Yeung et al. (2011). The treatment of Fulvetta follows Pasquet et al. (2006), while Moyle et al. (2012), and Gelang et al. (2009) were used to place the other small genera.

Leader et al. (2013) found that the Chinese Hill Warbler, Rhopophilus pekinensis, consists of two species. Accordingly, it has been split into Tarim Babbler, Rhopophilus albosuperciliaris, and Beijing Babbler, Rhopophilus pekinensis.

There seems to be insufficient reason to consider Northern Parrotbill, Paradoxornis polivanovi, as a separate species from Reed Parrotbill, Paradoxornis heudei, so they are merged as Reed Parrotbill, Paradoxornis heudei (see Penhallurick and Robson, 2009; Yeung et al., 2011).

The distinctive Yunnan Parrotbill, Sinosuthora ricketti, has been split from the Brown-winged Parrotbill, Sinosuthora brunnea. See Yeung et al. (2011) for genetic support and Wright et al. (2014) for photos.

Zosteropidae: White-eyes Bonaparte, 1853

13 genera, 142 species HBW-13

Zosterops itself is known as a “great speciator” for its propensity to invade new areas and diversify into new species and subspecies. (Diamond et al., 1976; Mayr and Diamond, 2001). Zosterops and its relatives have colonized and re-colonized the same islands and mountain ranges. Sometimes an island has been colonized by a single species that diversifies, sometimes an island has been invaded by various groups of white-eyes. This makes it hard or even impossible to sensibly group Zosteropidae species on geographic grounds. Although many of the clades make geographic sense, the colonization and re-colonization means the overall picture is hard to understand geographically unless you overlay geography on the phylogeny.

What is a White-eye?

What is a white-eye? Well, its a small bird with a prominent white eye ring. Of course, this is a naive definition and some members of the white-eye family (Zosteropidae) don't have eye-rings, while certain other small birds with white eye-rings are not part of Zosteropidae. More seriously, there are two parts to the question. What families are the closest relatives of the white-eyes, and what genera and species are part of the white-eye family.

Traditionally, the white-eyes were considered close to the honeyeaters (Meliphagidae, now part of Corvida), flowerpeckers (Dicaeidae, Passeroidea), and sunbirds (Nectariniidae, Passeroidea). Sibley and Ahlquist (1990) considered the white-eyes sylvioids. Cibois et al. (2003) went further, placing the Zosteropidae firmly in the Sylviidae/Timaliidae group. More recent analysis have confirmed this (e.g., Fregin et al., 2012). The white-eyes are sister to the three core babbler familes, Timaliidae, Pellorneidae, and Leiothrichidea.

There have also been changes in our notions of what species are part of the white-eye family. Cibois et al. (2003) noticed a close relationship between Zosterops and Yuhina (except for Yuhina zantholeuca, now placed in the Erpornis, in the Vireonidae). In combination with Cibois et al. (2002), this implied a close relation to a number of species then considered Stachyris babblers. These have been added to the Zosteropidae.

There have also been some surprising losses. Van Balen (2008) listed the Cinnamon Ibon, Hypocryptadius cinnamomeus, in a separate, monotypic, subfamily, but suggested it may not belong with the white-eyes at all. More recently, Moyle et al. (2009a) showed that it is not a white-eye, and suggested it may not even be a sylvioid. The final piece of this saga was the paper by Fjeldså et al. (2010). They found that the Cinnamon Ibon not only a member of Passeroidea, but is actually a passerid, a member of the Old World Sparrow family, Passeridae.

More recently, another species has been removed from the Zosteropidae. Alström et al. (2015a) found that the Madanga, formerly Madanga ruficollis (previously Heleia on TiF), is not a white-eye. It turns out that it is another member of Passeroidea, in fact, a pipit. It gets the new scientific name Anthus ruficollis. It's closest relative is the Alpine Pipit, Anthus gutturalis.

Basal Groups

The more recent work by Zhang et al. (2007) and Moyle et al. (2009a) has clarified how the yuhinas and former Stachyris babblers relate to the white-eyes, with Cibois et al. (2002), Cibois (2003), Collar and Robson (2007), and Moyle et al. (2009a) providing valuable information. The various Yuhinas are not monophyletic, but form a basal grade in Zosteropidae. Available evidence suggests they should be split into 3 or perhaps even 4 genera. Here hey are split into 3 genera, Yuhina, Staphida, and a currently unnamed genus, here designated “Yuhina”. The possible further split of brunneiceps and nigrimenta would also require a new genus name. The various Yuhina groups are comprised mostly of mainland Asian species (Staphida everetti occurs on Borneo).

After the yuhinas, the arrangement that seems to best fit the available studies is to put most of the remaining basal Zosteropidae into two main clades, with the Chestnut-faced Babbler, Zosterornis whiteheadi, separate from both. The first group seems to have two branches, a Philippine branch of former Stachyris babblers (Dasycrotapha and Sterrhoptilus) and a Micronesian branch. The latter includes the former honeyeater Apalopteron as well as some non-Zosterops white-eyes. Apalopteron and Cleptornis are closer to each other than to Zosterops (Springer et al., 1995), and may be sisters. The remaining Rukia may also be close to Cleptornis (the former Yap Rukia has moved to Zosterops and been given a new name, Olive-colored White-eye).

Then comes the Chestnut-faced Babbler, Zosterornis whiteheadi, on its own, with the last of the other basal Zosteropidae clades sister to Zosterops. The last basal group consists of the striped-babblers, another group of former Stachyris babblers from Philippines sometimes placed in Zosterornis. These lack a genus name and could also use a new English name. They are designated “Zosterornis” here. The other group includes the rest of the non-Zosterops white-eyes (a few have been merged into Zosterops). Since Madanga has been ejected, these would include the genera Tephrozosterops, Oculocincta, Lophozosterops, and Heleia, although I had separated some of the Lophozosterops into Apoia. Mayr (1965) argued that most of them, perhaps all, should be merged into Heleia. Mees (1969), disagreed and seemed to have won the point.

More recently, Moyle et al. (2009a) found that Zosterops wallacei was not part of Zosterops, but belonged near Lophozosterops superciliaris. Moreover, they found Lophozosterops to be paraphyletic with respect to Z. wallacei and Oculocincta. Gelang et al. (2009) found Lophozosterops paraphyletic with respect to Heleia. Taking all this into account, I think Mayr had the right idea and have placed all eleven species in Heleia.

Zosterops Overview

This brings us to Zosterops itself. It is a very large genus consisting of slightly over one hundred species. This diversification seems to be fairly recent, as befits a “great speciator”. Indeed, genetic investigations have revealed that Zosterops is an even greater speciator than we thought. If Cox's (2013) calibration is correct, this diversity has appeared in less than 4 million years.

The white-eyes propensity to speciate has also created a confused biogeographic picture. White-eyes have gone wild in invading new areas and forming new species. Sometimes, the same island has been invaded by more than one lineage in a short amount of time. Islands that seemed to be populated by two or more closely related white-eyes in a single invasion that then split are now known to have undergone two or more invasions. Nearby islands don't necessarily contain closely related white-eyes, even when they have been considered conspecific.

The treatment of Zosterops here draws on Slikas et al. (2000), Warren et al. (2006), van Balen (in HBW-13, del Hoyo et al., 2008), Moyle et al. (2009a), Melo et al. (2011), Cox (2013), and Cox et al. (2014). Cox et al., Warren et al., and Melo et al. focus coverage of African white-eyes. Compared to Warren et al., the closely related paper by Melo et al. (2011) increases sampling of the Gulf of Guinea species. Moyle et al. focus on the white-eyes of South-east Asia, Australasia, and Oceania. The genera Chlorocharis, Speirops and Woodfordia have been merged into Zosterops.

We still lack a good, complete phylogeny of the white-eyes. Moyle et al. (2009a) and Cox (2013) come closest. Moyle et al. focus on the eastern white-eyes, while Cox concentrates on the African white-eyes. Unfortunately, the basal portions of their phylogenetic trees are somewhat incongruent (e.g., where should the Silvereye Z. lateralis go?). This may be partially due to the rapid speciation the white-eyes have undergone, resulting in many short and poorly resolved internodes. I think the subclades are more reliable than their overall arrangement.

I have cobbled these together in a somewhat speculative fashion. This leaves Zosterops with three basal clades. There is an East Indian clade that is based on a fair amount of speculation, some traditional taxonomic views summed up by van Balen (2008), and DNA analysis of two of its species.

East Indian Clade: The hypothesized East Indian clade includes the atriceps and chloris superspecies. Given that some white-eyes thought to be closely related have turned out to be not so close, it wouldn't be surprising to find that some of the species tenetatively placed in this clade don't belong here. Indeed, the Ashy-bellied White-eye, Zosterops citrinella, was thought to belong to the chloris superspecies, but Moyle et al. (2009a) placed it elsewhere. I've included some species of uncertain affinities as basal members. Some of them are thought to be close to the atriceps clade.

It makes biogeographic sense to have the East Indian clade in a relatively basal position. The closest relatives to Zosterops are Heleia and “Zosterornis”. Both are restricted to the Malay Archipelago, with “Zosterornis” in the Philippines and Heleia primarily in Indonesia.

The other two clades are more solid, although support for them is still weak. There is an Australasian/Oceanic clade found by Moyle et al. (2009a), and there is the African clade studied in detail by Cox (2013).

Zosterops, African Clade

The African clade includes a basal group (the Oriental clade), followed by a number of African clades. The topology for the African species is based on Warren et al. (2006), Melo et al. (2011), Cox (2013), and Cox et al. (2014). The African clades are marked with letters A-F as in Cox (2013).

Oriental White-eyes: This clade includes the Japanese White-eye, Zosterops japonicus, and probably the Lowland White-eye, Zosterops meyeni. It also includes the Oriental White-eye, Zosterops palpebrosus, and its close cousin, the Sri Lanka White-eye, Zosterops ceylonensis. The range of the Oriental White-eye stretches from China to Arabia, providing a link between the eastern and African white-eyes.

Ancient Indian Ocean White-eyes (A): The first African white-eyes are the Ancient Indian Ocean White-eyes, Z. semiflavus through Z. mauritianus. These white-eyes occur on Maruritius, La Réunion, Grande Comore, and formerly the Seychelles (semiflavus). Warren et al. (2006) discovered that these white-eyes formed a distinct group, and that they were not closely related to the Recent Indian Ocean White-eyes (clade E3). In other words, there were two separate invasions of the the islands in the western Indian Ocean. This clade stems from the initial invasion.

Warren et al. (2006), Melo et al. (2011), and Cox et al. (2014) find this group to be quite basal, I think too much so to make biogeographic sense. I have followed Cox (2013) on this where it is merely the most basal African clade. The time calibrated phylogeny in Cox et al. (2014) suggests that this “ancient” clade may be only 2 million years old. Still, it is ancient compared to the 1 million year ago origin of clade (E3), the Recent Indian Ocean white-eyes.

Socotra White-eye (B): This is an interesting case. Kirwan (2007) argued that the 4 white-breasted subspecies of abyssinicus are closely related. Indeed, he even argued that socotranus and abyssinicus are virtually indistinguishable. Cox's (2013) results upset that conclusion. However, Cox found that the Socotran socotranus ended up on a separate branch from the mainland socotranus. The mainland socotranus group with abyssinicus. As a result, I'm treating the Socotran socotranus as a separate species, Socotra White-eye, Zosterops socotranus.

Middle African White-eyes (F): The restructing of the Montane and African Yellow White-eye is on display in this clade, which ranges across the mid-section of Africa from Nigeria and the Gulf of Guinea to Pemba Island (near Zanzibar). It contains 3 pieces of lowland and montane white-eyes mixed together with a couple of Speirops and Pemba White-eye (sometimes included in African Yellow). We start with a pair of highland taxa that appear to be sister, the Mount Cameron Speirops, Zosterops melanocephalus, and the newly split Kilimanjaro White-eye, Zosterops eurycricotus (from Montane White-eye, Zosterops poliogastrus). Next up is the Albertine White-eye, Zosterops stuhlmanni. This is a rebranded version of the previously split Kivu White-eye, which has added stuhlmanni and toroensis. As stuhlmanni has priority, the scientific name had to change, and since the range is now larger, encompassing a big chunk of the Albertine rift area, it also got a new English name. It also contains the Forest White-eye, Zosterops stenocricotus, which has been split from African Yellow White-eye, Zosterops senegalensis (see also Melo et al, 2011).

Abyssinian White-eye (D): These species are followed by the Abyssinian White-eye, Zosterops abyssinicus, represented in Cox et al. (2014) by socotranus. Kirwan (2007) argues that the 4 white-breasted subspecies of abyssinicus are closely related. Indeed, he argues that socotranus and abyssinicus are virtually indistinguishable. The other two are arabs, which is slightly different, and omoensis, which is more easily distinguishable. Indeed, Cox (2013) groups the mainland white-breasted abyssinicus together. Cox (2013) found that the yellow-breasted subspecies flavilateralis and jubaensis (Kenya White-eye, Z. flavilateralis of clade E1) are not closely related to mainland socotranus, and the entire white-breasted group.

Gulf of Guinea White-eyes (C): The Gulf of Guinea White-eyes include the other two former Speirops (Principe and Black-capped), along with the Principe White-eye, Zosterops ficedulinus, Annobon White-eye, Zosterops griseovirescens, and Sao Tome White-eye, Zosterops feae. Note that the last has been split from Z. ficedulinus. One result of all this is that the Gulf of Guinea white-eyes result from two separate expansions.

Southern African White-eyes (E2): The Southern African White-eyes include two additional splits: South Pare White-eye, Zosterops winifredae, from Z. poliogastrus, and Southern Yellow White-eye, Zosterops anderssoni, from Z. senegalensis. I had previously use the name Nyasa White-eye, Zosterops stierlingi, but the extensive taxon sampling in Cox (2013) has changed the picture. The species clade includes not just Zosterops stierlingi, but also tongensis and at least some portion of anderssoni (samples from Angola and Zambia). This likely means the type of anderssoni (from Nambia) is also included here, so use of the name anderssoni is justified. A sample of anderssoni from Malawi grouped with the Cape White-eye. The one sample of quanzae showed up on an earlier branch, together with a couple of samples from heinrichi. Oddly, samples of heinrichi from the same location ended up grouped with the Cape White-eye. I suspect that further work will clarify where these belong. Cox (2013) also sampled kasaicus, but did not include it in any of the trees. According to the text, the samples ended up somewhere in the winifredae-virens group.

East African Clade (E1): Cox (2013) and Cox et al. (2014) have two species from East Africa next, the yellow-breasted `abyssinicus', Kenya White-eye, Z. flavilateralis (including jubaensis) and its sister taxon Z. poliogastrus mbuluensis, which I've promoted to species rank as Mbulu White-eye. The treatment mbuluensis as a species is also supported by Meimberg et al. (2016). Thus we maintain the traditional distinction between the highland (mbuluensis) and lowland (flavilateralis) white-eyes, but in a new form. Note that many of the highland forms in the range of flavilateralis are not so closely related to it, but that mbuluensis is.

Recent Indian Ocean Clade (E3): The arrangement of these species is based on Warren et al. (2006), who sampled more genes than Cox (2013). There are still some questions concerning species boundaries within the maderaspatanus group of Indian Ocean white-eyes. I've split Anjouan White-eye, Zosterops anjuanensis, from Malagasy White-eye, Zosterops maderaspatanus, on the grounds that it is more distantly related to Z. maderaspatanus than either Z. kirki or Z. mayottensis. It has historically been called “Lesser White-eye”. I've also split Moheli White-eye, Zosterops comorensis, from Mayotte White-eye, Zosterops mayottensis (or in two steps from Malagasy White-eye, Zosterops maderaspatanus). The three taxa (maderaspatanus, comorensis, mayottensis) seem about equally distant. For similar reasons, I've also separated Aldabra White-eye, Zosterops aldabrensis, from Kirk's White-eye, Zosterops kirki (or in two steps from Malagasy White-eye, Zosterops maderaspatanus).

As far as the allocation of subspecies within the maderaspatanus clade of Indian Ocean White-eyes is concerned, the extinct Seychelles Yellow White-eye, Z. semiflavus, often considered a subspecies of Z. mayottensis, is treated as a separate species because it is not even part of this group, instead belonging to the “ancient Indian Ocean White-eyes”. This leaves voeltzkowi and likely menaiensis as subspecies of Z. maderaspatanus.

Current Division of Traditional White-eyes
(compared to HBW vol. 13)
The headings indicate species in HBW-13. Below them are the allocation of subspecies to the species on the TiF list. The subspecies with question marks indicate either a lack of genetic testing (menaiensis), or a subspecies that was tested but not placed in the tree (kasaicus). The subspecies in red appeared in two different groups in Cox (2013) (anderssoni, gerhardi, and heinrichi). Meimberg et al. (2016) also supports species status for kikuyuensis, mbuluensis, and silvanus. Although they claim support for species status for kulalensis, it is certainly not evident from their genetic analysis, with only a single sample closely allied to poliogastrus.

Abyssinian White-eye, Zosterops abyssinicus
Socotra White-eye
Z. socotranus (A)
socotranus (Socotra)
Abyssinian White-eye
Z. abyssinicus (D)
abyssinicus, arabs, socotranus (mainland),
omoensis
Kenya White-eye
Z. flavilateralis (E1)
jubaensis, flavilateralis

Cape White-eye, Zosterops pallidus
Orange River White-eye
Z. pallidus (E2)
pallidus, sundevalli
Cape White-eye
Z. virens (E2)
capensis, caniviridis, virens, atmorii,
heinrichi, anderssoni, kasaicus??

African Yellow White-eye, Zosterops senegalensis
Albertine White-eye
Z. stuhlmanni (F)
toroensis, reichenowi, stuhlmanni
Forest White-eye
Z. stenocricotus (F)
stenocricotus
Northern Yellow White-eye
Z. senegalensis (F)
senegalensis, demeryi, gerhardi, jacksoni
Southern Yellow White-eye
Z. anderssoni (E2)
stierlingi, kasaicus??, heinrichi, quanzae,
anderssoni, tongensis

Montane White-eye, Zosterops poliogastrus
Kilimanjaro White-eye
Z. eurycricotus (F)
eurycricotus
Kikuyu White-eye
Z. kikuyuensis (F)
kikuyuensis
Montane White-eye
Z. poliogastrus (F)
poliogastrus, kaffensis, kulalensis, gerhardi
Mbulu White-eye
Z. mbuluensis (E1)
mbuluensis
Taita White-eye
Z. silvanus (E2)
silvanus
South Pare White-eye
Z. winifredae (E2)
winifredae, kasaicus??

Malagasy White-eye, Zosterops maderaspatanus
Anjouan White-eye
Z. anjuanensis (E3)
anjuanensis
Aldabra White-eye,
Z. aldabrensis (E3)
aldabrensis, menaiensis??
Kirk's White-eye
Z. kirki (E3)
kirki
Malagasy White-eye
Z. maderaspatanus (E3)
menaiensis??, maderaspatanus, voeltzkowi
Moheli White-eye
Z. comorensis (E3)
comorensis

Letters in parentheses refer to clades in the tree.

Zosterops, Australasian/Oceanic Clade

The Australasian/Oceanic Clade includes three subclades: a western Australasian clade, an eastern Australasian clade, and a Micronesian/Melanesian clade.

East Asian/Oceanic Clade: This clade includes white-eyes from China, the Malay Archipelago, and western Micronesia (Rota and Yap). The second subclade contains the Yellowish White-eye, Zosterops nigrorum. This brings us to the last portion of Zosterops, the 3-part clade. The first subclade may actually be basal in Zosterops, with all other clades being subclades. However, Moyle et al. (2009a) prefer a different topology which is followed here. They found that the Mountain Blackeye, Chlorocharis emiliae, is actually a Zosterops, apparently closely related to the Mountain White-eye. The placement of the Rota White-eye, Zosterops rotensis, and the Plain (formerly Yap) White-eye, Zosterops hypolais is inspired by Slikas et al. (2000).

West Australasian Clade: I've split one species in the western clade. The two representatives of Z. palpebrosus were widely separated, one in this clade, one in clade II. I have promoted the one here (Z. palpebrosus melanurus) to species status. Although it has apparently been considered a separate species in the past (HBW13), I couldn't find an English name. I'm temporarily designating it “Sunda White-eye” on geographic grounds. When considered a separate species, melanurus has contained two subspecies: melanurus and unicus. The subspecies unicus is what Moyle et al. actually sampled. As buxtoni is known to hybridize with melanurus, and palpebrosus is rather distant genetically, it is provisionally treated as a subspecies of melanurus, even though it is visually distinct. The other races of Oriental White-eye remain with palpebrosus. This means the highland race in Sumatra (buxtoni) is in the melanurus group, while the lowland race (auriventer) belongs to palpebrosus.

This clade also include the Ashy-bellied White-eye, Zosterops citrinella, which had been considered to possibly belong to the chloris superspecies. It doesn't, unless the results of Moyle et al. (2009a) are highly misleading.

East Australasian Clade: The second subclade includes the griseotinctus superspecies of the Solomons and nearby islands (minus the Banded White-eye, Zosterops vellalavella, which does not really belong with the others) and the Lifou White-eyes, which may be sister species (Cornetti et al., 2015). It also contains the Silvereye, Zosterops lateralis, and closely related species. However, even without vellalavella, the rest of the griseotinctus group is still not monophyletic, with some closer to the Silvereye than to other supposed group members.

The Dark-eyed White-eye, Zosterops tetiparius, including paradoxus (sometimes called rendovae) has been split from the Solomons White-eye, Zosterops kulambangrae, based on H&M-4, IOC 6.2, and Moyle et al. (2009).

The rendovae problem

There is a long-running controversy over the name Zosterops rendovae. The origin of the problem dates back to the 1880's. Ramsay (1881) named a new species of white-eye from San Cristobal (Makira) Island as Tephras olivaceus. Tristram (1882) moved the species to Zosterops. This meant that it needed a new name as there already was a Zosterops olivaceus. Tristram named it rendovae and included both a statement that it applied to Ramsay's bird and a description. Unfortunately, the description was not a description of Ramsay's bird, but of a white-eye from Rendova Island, not San Cristobal. In other words, Tristram had unwittingly applied the name to two taxa at once. Later that same year, Ramsay (1882) provided another replacement name (ugiensis) for T. olivaceus. Still later that same year, Salvadori recognized the double definition by Tristram. He restricted the name rendovae to the Rendova island white-eyes and proposed another replacement, ramsayi, for T. olivaceus. As it happens, Z. ramsayi was both preoccupied (by Z. ramsayi Masters 1876) and a junior synonym of the existing names. Some confusion about the San Cristobal specimen led to two more names being applied to the white-eyes there (salomonensis and alberti), but they need not concern us further.

Salvadori's designation of the Rendova white-eyes as rendovae was generally accepted and until 1955, although there was some confusion about how many white-eyes were on San Cristobal (and which name to use). Eventually it became clear there was only one and that ugiensis was the oldest available name. That would make the Solomons White-eye, Zosterops rendovae rather than Zosterops kulambangrae as it includes the Rendova population.

Mees (1955) argued that Salvadori's description was irrelevant for a replacement name, and that rendovae properly applied to T. olivaceus. He coined a new name (paradoxus) for the Rendova population. In this case, the Gray-throated White-eye would be Zosterops rendovae instead of Zosterops ugiensis. This solution has also been adopted by some. Others, such as Galbraith (1957), have insisted that Salvadori is correctly regarded as first reviser. This conflict has created considerable confusion concerning rendovae.

There had been a recent trend to suppress rendovae, although I do not believe that it has been made official. Clements does not use it. Recently, both H&M-4 and IOC6.2 have brought it back. In doing so, they have illustrated the problem with using rendovae. For H&M-4, Z. rendovae is the Dark-eyed White-eye, consisting of subspecies rendovae = paradoxus and tetiparius. For IOC 6.2, Z. rendovae is the Gray-throated White-eye, with subspecies rendovae = ugiensis, oblitus, and hamlini. In other words, H&M-4 and IOC 6.2 have provided the perfect illustration of the rendovae problem.

Although I think H&M-4 are technically correct, regarding Salvadori as first reviser, I nonetheless continue to treat rendovae as supressed. Using it is an impediment to accurate communication. Thus I call the Rendova white-eyes Zosterops tetiparius paradoxus and the San Cristobal white-eyes Zosterops ugiensis ugiensis. The Dark-eyed White-eye is then Z. tetiparius and the Gray-throated White-eye is Z. ugiensis.

NB: The Biodiversity Heritage Library provides access to the relevant original papers.

Micronesian/Melanesian Clade: Moyle et al. also included two subspecies of Z. ugiensis (ugiensis and hamlini) in their analysis. They ended up in different parts of the Micronesian/Melanesian subclade, with ugiensis itself more closely related to the Micronesian branch. There does not appear to be an English name for Z. hamlini. Since the hamlini subspecies is restricted to Bougainville Island, I'm referring to it as the “Bougainville White-eye”. The other subspecies, Z. ugiensis oblitus, is restricted to Guadalcanal. Given the proximity of Guadalcanal to San Cristobal (home of Z. u. ugiensis, I'm treating it a subspecies of Z. ugiensis for the present.

It's not clear exactly where to put the Bridled White-eye, Zosterops conspicillatus. Slikas et al. (2000) argued that it was not conspecific with Zosterops semperi. However, some trees put it in a clade with Z. semperi and the Olive-colored White-eye, Zosterops oleagineus (formerly in Rukia). Moyle et al. (2009a) included Z. cinereus in a clade with those two and Z. ugiensis, but didn't analyze Z. conspicillatus. I've bundled them all together with the other member of the cinereus superspecies, Z. finschii.

Based on Hayes et al. (2016), the Gray-brown White-eye, Zosterops cinereus, has been split into Kosrae White-eye, Zosterops cinereus, and Gray-brown White-eye, Zosterops ponapensis.

The newly recognized Vanikoro White-eye, Zosterops gibbsi, (Dutson, 2008) is thought to be close to sanctaecrucis, which is probably near metcalfii and stressmani. Together, they are the other branch of the subclade.

Timaliidae: Babblers, Scimitar-Babblers Bonaparte, 1840

10 genera, 52 species HBW-12

Timaliidae is often attributed to Vigors & Horsfield, 1827 (e.g. Bock, 1994), but this is incorrect (Olson, 1995). It is attributed to Bonaparte by Brodkorb (1978).

Click for Timaliidae tree
Click for Timaliidae tree

Following Collar and Robson (2007), I had split Stachyridopsis from Stachyris. However, Moyle et al. (2012) found that the Chestnut-winged Babbler (formerly Stachyris erythroptera). also belongs in this group. It is the type species of Cyanoderma (Salvadori, 1874). This takes precendence over Stachyridopsis (Oates, 1883), and I have changed the names accordingly.

Moyle et al. also found that the Pin-striped Tit-Babbler is more closely related to Timalia than to the remaining Macronus species. It is the type of Mixornis (Blyth, 1842). It seems likely that kelleyi, bornensis (split from gularis by Collar, 2006), and flavicollis are close relatives. The other two Macronus are a bit more distant (Moyle et al., 2012).

Gelang et al. (2009) showed that Xiphirhynchus is embedded in Pomatorhinus, so it has been merged into Pomatorhinus here. More recently, Dong et al. (2010) showed that Pomatorhinus should be split into two genera. The name Erythrogenys (Baker 1930, type erythrogenys) applies to the them. This group is sister to the Pomatorhinus-Stachyris clade. Finally, Moyle et al. also found that Sphenocichla is embedded in Stachyris.

The scimitar-babblers (Pomatorhinus and Erythrogenys) have been the subject of several detailed studies, including Dong et al. (2010, 2014), Nyári and Reddy (2013), and Reddy et al. (2011). It is likely that some of the existing species will be split, but the best way to do that is not yet clear. What is clear is that species limits need to be changed for the Streak-breasted Scimitar-Babbler, Pomatorhinus ruficollis, Chestnut-backed Scimitar-Babbler, Pomatorhinus montanus, and White-browed Scimitar-Babbler, Pomatorhinus schisticeps. For now, the White-browed Scimitar-Babbler (P. schisticeps) loses all its subspecies that lie east of the Irrawaddy River. Three, nuchalis, fastidiosus, and klossi join the Chestnut-backed Scimitar-Babbler (P. montanus), while the olivaceus group (olivaceus, ripponi, difficilis, humilis, and annamensis) joins the Streak-breasted Scimitar-Babbler, P. ruficollis. Keep in mind that these are subject to further splits.

Pellorneidae: Fulvettas, Ground Babblers Delacour, 1946

14 genera, 60 species Not HBW Family

Click for Pellorneidae tree
Click for Pellorneidae tree

I've characterized Pellorneidae as ground babblers because so many are ground and understory feeders, moreso than the babblers not part of Pellorneidae.

The genus Alcippe is another poster child for the DNA revolution. The former Alcippe has been broken into four parts, with the Bush Blackcap ending up in Sylvia (Sylviidae) via Lioptilus, the Fulvetta fulvettas in Paradoxornithidae, the Schoeniparus fulvettas (including Pseudominla) here in Pellorneidae, and the remaining Alcippe somewhat tentatively placed basally in Leiothrichidae. In fact, there's still considerable uncertainty about the status of Alcippe proper. Various genes place it basally in Pellorneidae, while others place it basally in Leiothrichidae. Moyle et al. (2012) grappled with this issue, finally settling on Leiothrichidae, which I follow for now.

Since Moyle et al. (2012) have very good taxon sampling for most genera, I mostly follow their preferred arrangement. However, they include neither Graminicola nor Turdinus. Fortunately, Gelang et al. (2009) did include them. They found them basal to the remaining Pellorneidae. All of Turdinus has been included in Napothera. Collar (2006) argued for separating them in Turdinus (see also Collar and Robson, 2007). The one species of Turdinus sampled by Gelang et al. proved quite different from Napothera.

Working down the tree, Ophrydornis has been returned to Malacopteron. It is sister to Gampsorhynchus-Schoeniparus. Note that Schoeniparus, which was previously split from Alcippe (Pasquet et al., 2006; Collar and Robson, 2007), includes Pseudominla. The remaining portion of Pellorneidae is a well-supported clade in Moyle et al. (2012). Although Illadopsis seems to branch off next, there is evidently some remaining uncertainty about whether that is correct, and how the pieces of this clade fit together.

It's clear that the species in Illadopsis belong together (once Ptyrticus has been absorbed), as do those in the expanded Pellorneum. This version of Pellorneum has absorbed Trichastoma and a couple of species from Malacocincla. After Kenopia, there are two well-supported clades. One contains the remaining Malacocincla and the three Gypsophila (once part of Napothera). The other contains Ptilocichla, the one remaining Napothera species, and Rimator. Moyle et al. recommend merging Rimator and Napothera, but I prefer to leave the distinctive Rimator group in its own genus. Note that Rimator includes Jabouilleia.

The genera Leonardina and Robsonius are no longer part of this family. Indeed, they are not even part of the larger babbler clade. Oliveros et al. (2012) found that Leonardina belongs to the Old World flycatcher family Muscicapidae and that Robsonius is in the grassbird family Locustellidae.

The Pale-breasted Thrush-Babbler, Illadopsis rufipennis may involve more than one species. Nguembock et al. (2009b) found samples of I. rufipennis extrema basal to a clade containing pyrrhoptera and other rufipennis.

The Rufous-rumped Grassbird, Graminicola bengalensis, has been split into Chinese Grassbird, Graminicola striatus, and Indian Grassbird, Graminicola bengalensis (Leader et al., 2010). As the English name suggests, the Chinese Grassbird includes the race sinicus.

Olsson et al. (2013) found that the Rufous-vented Prinia, Laticilla burnesii does not belong in Prinia. It has been placed in Laticilla (Blyth 1845, type burnesii) and moved to the family Pellorneidae. The Swamp Prinia is thought to be a close relative, so it has become Laticilla cinerascens. Price et al. (2014) found L. burnesii closer to Napothera and Rimator than to Malacocincla. The primary English name of these two species has been changed from Prinia to Grass-Babbler (Rufous-vented Grass-Babbler and Swamp Grass-Babbler).

Leiothrichidae: Laughingthrushes Swainson, 1831

27 genera, 145 species Not HBW Family

Click for Leiothrichidae tree
Click for Leiothrichidae tree

The final piece of the sylviid/babbler clade is Leiothrichidae, the laughingthrush family. I had previously treated it as two clades ranked as subfamilies based on Gelang et al. (2009) and especially Luo et al. (2009). However, neither included the Striated Laughingthrush, Grammatoptila striata, and both placed Alcippe outside Leiothrichidae. Moyle et al. (2012) included both taxa and found that they, and probably Cutia take the basal positions in Leiothrichidae, precluding a simple two-clade structure.

As mentioned previously, the Moyle et al. topology puts the Alcippe fulvettas a as basal group in Leiothrichidae. The Gray-cheeked Fulvetta, Alcippe morrisonia has been split into several species based on Zou et al. (2007) and Song et al. (2009).

Garrulax itself has often been treated as a large genus. However, that doesn't work well as parts of it don't belong together. Before adequate genetic testing had been done, Collar and Robson broke Garrulax into 11 genera: Dryonastes, Melanocichla, Rhinocichla, Pterorhinus, Grammatoptila, Stactocichla, Leucodioptron, Strophocincla, Ianthocincla, and Trochalopteron, and Garrulax. They also grouped Babax with the laughingthrushes. We retain about half of these genera.

The monotypic Grammatoptila takes a relatively basal position in Leiothrichidae, away from the other laughingthrushes. Cutia is the next branch. The family then divides in two. The first clade include Turdoides and Garrulax. The other includes the Trochalopteron laughingthrushes and a number of smaller genera including the barwings, minlas, sibias, and other species.

If you examine the species-level tree, you will see many question marks indicating untested species and many polytomies. Obviously, some further revision will be needed.

The Turdoides group is sister to the main laughingthrush clade (containing Garrulax). Moyle et al. (2012) found that Phyllanthus and Kupeornis are embedded in it and this had previously led me to use a broad Turdoides. This has been rearranged and divided taking Price et al. (2014) into account. The two embedded genera have been restored, and I have also separated Acanthoptila, Argya, Chatarrhaea, and Malacocircus from Turdoides. This result is similar to H&M-4 (Dickinson and Christidis, 2014) and the HBW BirdLife Checklist (del Hoyo and Collar, 2016), except that three species have been placed in Malacocircus based on Price et al. (2014).

The Garrulax group has been divided into six genera (one uncertainly placed): Leucodioptron, Stactocichla, Garrulax, Melanocichla, Ianthocincla, and Pterorhinus. Various papers give various results for the arrangement of Pterorhinus. I have based the structure of Pterorhinus on Päckert et al. (2011), with assistance from Luo et al. (2009), Moyle et al. (2012), Robin et al. (2017), and Wu et al. (2014).

The other big piece of Leiothrichidae contains the Trochalopteron laughingthrushes. Based on Robin et al. (2017), the Kerala Laughingthrush, Trochalopteron fairbanki, and Black-chinned Laughingthrush, Trochalopteron cachinnans, have been moved to Montecincla. Further, there are splits:

Both Trochalopteron and Montecincla are basal to a group that includes Heterophasia sibias (including Malacias), Leiothrix (including Mesia), Minla (including Leioptila), Laniellus (formerly called Crocias, see below), Liocichla, followed by the Actinodura group including Sibia, Siva, and Chrysominla. Some of the arrangement of species is based on Dong et al. (2010).

Following Gregory and Dickinson (2012), the genus name Laniellus Swainson, 1832 has replaced Crocias Temminck, 1836.

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