Sylvioidea III

Passerines

Tyranni: Suboscines

Passeri: Oscines

Passerida

Sylvioidea
Muscicapoidea and allies
Passeroidea

The 45 Orders

Paleognaths

Galloanserae

Metaves

Pelecanae

Charadriae

Passerae

Babblers and allies

Timaliidae 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 Gelang et al. (2009), which I interpret as evidence in favor of grouping the remaining 450 babblers and allies into 5 families.

One is the new Sylviidae, which includes the parrotbills. The white-eyes get their own family, Zosteropidae. Then comes a more narrowly circumscribed Timaliidae, followed by the Pellorneidae. It ends with the laughingthrush family, Leiothrichidae.

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. None of these names appear to have had much recent use, and priority between them seemed unresolved. Because of this, Gelang et al. could able to abandon Leiothrichidae (for Turdoididae), but they have endorsed it (as Leiothrichinae) instead, and I follow their usage.

Sylviidae: Sylviid Warblers, Parrotbills, Fulvettas Leach, 1820

18 genera, 70 species HBW-11

Although greatly reduced, the Sylviidae are still a respectable family containing nearly 70 species. It includes Sylvia itself, together with a few of the old Sylviidae, some babblers, and a former cisticolid (Rhopophilus). Although some uncertainty remains, we now have a good-quality species-level tree for the Sylviidae based mainly on the work of Gelang et al. (2009), Pasquet et al. (2006), Voelker and Light (2011), and Yueng et al. (2011).

Click for Sylviidae tree
Click for Sylviidae tree

The Lioparus and Fulvetta fulvettas had previously been in the genus Alcippe (Timaliidae). The order here relies on Cibois (2003), Pasquet et al. (2006), and Gelang et al. (2009). In the former two, the Sylviidae are clade 2. Pasquet et al. recommend the use of Lioparus and Fulvetta, and Collar and Robson (2007) helped clarify the limits of those genera.

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 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. Accordingly, I've submerged Horizonrhinus, Lioptilus, and Pseudoalcippe into Sylvia. The Abyssinian Catbird, Parophasma galinieri, belongs in the clade containing Sylvia, but its exact position is also uncertain (Gelang et al., 2009). I'm guessing that it belongs close to Sylvia also (near Curruca is the other possibility).

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. There remains some uncertainty about the relationships between the first 3 genera, but they are close and perhaps intertwined. Given that, it might make sense to merge them into one genus, Sylvia. The order within Curruca is based on Shirihai et al. (2001), Böhning-Gaese et al. (2003), and Böhning-Gaese et al. (2006). Exactly where Moupinia goes is not clear.

Finally, Myzornis has been moved here from incertae sedis, and belongs in an unresolved basal trichotomy (Gelang et al., 2009).

The arrangement within Curruca is based on Voelker and Light (2011). There are several notable clades which are similar to those described by Shirihai et al. (2001). Curruca nana and deserti could be called subgenus Atraphornis. Atraphornis is sister to the rest of Curruca, which divides into two major groups. In first group, C. nisoria (subgenus Adophoneus) is sister to the rest, which then divides into two clades. Subgenus Parisoma includes layardi, subcaerulea and boehmi, while curruca through leucomelaena are subgenus Curruca. The second group is subgenus Melizophilus, comprised of C. communis and the rest of genus Curruca.

Moltoni's Warbler, Curruca moltonii, (also known as subalpina, see Baccetti et al., 2007) was split from C. cantillans by Brambilla et al. (2008).

The phylogeny of the parrotbill-fulvetta group is not entirely certain. It is clear that there are 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 arrangment than the mitochondrial genes. The version here is a compromise. It 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, is Yeung et al. (2011). The treatment of Fulvetta follows Pasquet et al. (2006), while Gelang et al. (2009) was used to place the other small genera.

Zosteropidae: White-eyes Bonaparte, 1853

19 genera, 136 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.

The position and composition of the Zosteropidae has recently undergone some changes. Traditionally, they were considered close to the Meliphagidae (now part of Corvida), Dicaeidae, and Nectariniidae (both now part of Passerida). Sibley and Ahlquist (1990) considered them sylvioids. Cibois et al. (2003) went further, placing the Zosteropidae firmly in the Sylviidae/Timaliidae group. They also 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.

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. They are split into 4 genera, Yuhina, Staphida, and two currently unnamed genera, here designated Yuhina1 and Yuhina2. The various Yuhina groups are comprised mostly of mainland Asian species (Staphida everetti occurs on Borneo).

After the yuhinas, the other basal Zosteropidae seem to form a clade with three major subclades. One consists of the Zosterornis babblers of the Philippines (formerly Stachyris). The second subclade seems to have two branches, a Philippine Stachyris branch (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).

The third subclade includes the rest of the non-Zosterops white-eyes (a few have been merged into Zosterops). Moyle et al. (2009a) found that Zosterops wallacei was not part of Zosterops, but belonged near Lophozosterops superciliaris. There isn't another genus name available, so I've listed it as “Zosterops” wallacei. Gelang et al. (2009) also place Heleia here. These species from the Lesser Sundas form one branch of the third subclade. The other Lophozosterops seem to be in a sister branch, along with Oculocincta. As a result, the other Lophozosterops need a new genus name, and Apoia (type goodfellowi) is available (not Oreosterops, where Bonaparte's designation of montanus as type has priority). Madanga and Tephrozosterops may also belong to this Indonesian/Philippine subclade.

Zosterops Overview

This brings us to Zosterops itself. It is a very large genus consisting of nearly 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. Some islands that seemed to be populated by two or more closely related white-eyes in a single invasion that then split have actually 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), and Melo et al. (2011). Warren et al. have the most extensive coverage of African white-eyes. 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.

Moyle et al. (2009a) found a division into two main clades. Due to space considerations, each main clade gets a separate tree page. Neither clade is that well-supported, with clade I having stronger support. There is a possibility that clade I may be embedded in clade II. Nonetheless, I follow the best current estimate here, that of Moyle et al.

Zosterops, clade I

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

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). 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.

There is a long-running controversy over the name Zosterops rendovae. Since at least the 1950's, it has been applied to two different subspecies, one of which is in this subclades. I'm not clear on what the correct scientific name is here, but I use the same names as IOC. This translates to calling one of the subspecies Zosterops kulambangrae paradoxus (if it were rendovae, the species would also be rendovae) and the other one Zosterops ugiensis ugiensis. The three subspecies of Z. kulambangrae form a clade in Moyle et al. (2009a), and are treated here as a single species in the eastern Australasian clade. The other two subspecies are sometimes separated as Rendova White-eye, Zosterops paradoxus, and Tetepare White-eye, Zosterops tetiparius.

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.

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.

Zosterops, clade II

Clade II includes four parts: an Australasian clade and a 3-part clade stretching from east Asia to west Africa. It breaks into an Asian clade, the Yellowish White-eye, all by itself, and an African clade.

Australasian Clade: The Australasian clade includes the atriceps and chloris superspecies. I've included some species of uncertain affinities as basal members. Some of them are thought to be close to the atriceps clade. Only two of these species have been sequenced, and a lot of uncertainty remains here.

Asian Clade: 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).

Yellowish White-eye: The second subclade contains only a single species, the Yellowish White-eye, Zosterops nigrorum. It's not entirely clear where this Philippine species belongs. Although it seems to have some African affinities, it doesn't make sense to bury it in the African subclade.

African Clade: The last subclade includes two basal non-African members: the Oriental White-eye, Zosterops palpebrosus, and its close cousin, the Sri Lanka White-eye, Zosterops ceylonensis. The “ancient Indian Ocean” white-eyes of Warren et al. (2006) branch off next. Presumably Madagascar and nearby islands (Comoros, Réunion, Seychelles), were colonized by white-eyes as or before they arrived in Africa. Interestingly, there's a second clade of Indian Ocean white-eyes that originated after the white-eyes were well-established in Africa.

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”.

As far as the allocation of subspecies within the maderaspatanus clade 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. The race aldabrensis is grouped with Z. kirki, and comorensis with Z. mayottensis. Both should be considered for elevation to species level. This leaves voeltzkowi and menaiensis as subspecies of Z. maderaspatanus.

The topology for the African species is based on Warren et al. (2006) and Melo et al. (2011). Unlike Moyle et al., they have Z. abyssinicus sister to Z. poliogastrus mbuluensis, which I promote to species rank. Another African group comes next. It contains a problematic taxon, Z. kikuyuensis. The gene tree suggests that kikuyuensis from Mt. Kenya is not the closest relative of kikuyuensis from the Aberdares range. This could be a sampling problem, indicate interbreeding or introgression, or could represent a previously unknown species or subspecies.

A similar problem affects the Kenya White-eye, Zosterops flavilateralis (including Z. abyssinicus omoensis), which I treat as sister to a clade containing the new Indian Ocean white-eyes and a group of white-eyes from eastern and southern Africa. This group contains two other splits: South Pare White-eye, Zosterops winifredae, from Z. poliogastrus, and Nyasa Green White-eye, Zosterops stierlingi, from Z. senegalensis. This whole group, including Z. flavilateralis, is closely related to Speirops, which has been merged into Zosterops.

Melo et al. (2011) found that the former Speirops species are not a clade. Two of them group with the Forest White-eye, Zosterops stenocricotus, now split from African Yellow White-eye, Zosterops senegalensis. The other two group with 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.

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 may not even be a sylvioid. The latest news is from Fjeldså et al. (2010). They found that the Cinnamon Ibon is actually a passerid!

Timaliidae: Babblers, Tit-Babblers, Scimitar-Babblers Vigors & Horsfield, 1827

11 genera, 52 species HBW-12

Timaliidae tree The split of split Stachyridopsis from Stachyris follows Collar and Robson (2007). 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, the name is apparently based on an error of Hodgson's in 1836) applies to the remainder, which are sister to the Pomatorhinus-Stachyris clade.

Pellorneidae: Fulvettas, Ground Babblers Delacour, 1946

19 genera, 70 species Not HBW Family

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 portion of Alcippe in Pellorneidae has been split into 2 genera: Alcippe, and Schoeniparus. This follows Pasquet et al., (2006) rather than Collar and Robson (2007) who suggest 3 genera. Their Pseudominla is considered part of Schoeniparus here (contrary to some earlier versions of this list).

Many of the generic limits follow Collar and Robson (2007). Thus Jabouilleia joins Rimator, Ophrydornis is split from Malacopteron, and Robsonius, Turdinus, and Gypsophila split from Napothera.

The organization here draws on Pasquet et al. (2006), Collar and Robson (2007), Gelang et al. (2009). As you can see from the tree, information is lacking on several genera believed to be in Pellorneidae.

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 Gray-cheeked Fulvetta, Alcippe morrisonia has been split into several species based on Zou et al. (2007) and Song et al. (2009).

Leiothrichidae: Laughingthrushes Swainson, 1831

21 genera, 133 species Not HBW Family

The final piece of the sylviid/babbler clade is Leiothrichidae, the laughingthrush family. The analysis of Luo et al. (2009) suggests it contains two main clades, which we rank as subfamilies: Turdoidinae and Leiothrichinae.

Garrulax itself has often been treated as a large genus. However, that doesn't work well as parts of it end up in each tribe. The generic limits here are based on Collar and Robson (2007), but interpreted in light of Luo et al. (2009).

Collar and Robson broke Garrulax into 11 genera: Dryonastes, Melanocichla, Rhinocichla, Pterorhinus, Grammatoptila, Stactocichla, Leucodioptron, Strophocincla, Ianthocincla, and Trochalopteron, and Garrulax. This is too much. Luo et al. (2009) made clear that Dryonastes and Pterorhinus are not monophyletic. For now, it seems best to return most of the species in these two genera to Garrulax. Moreover, Babax should join them. We can usefully retain Ianthocincla and Leucodioptron, which together are sister to this version of Garrulax. These form the core of Turdoidinae. I'm not really sure where Melanocichla, Rhinocichla, Stactocichla, or Grammatoptila go, and have temporarily left them at the end of the Turdoidinae.

Leiothrichinae genera Leiothrichinae also contains a big chunk of the former Garrulax. Here Strophocincla seems to be nested within Trochalopteron, so I have merged them. It also seemed best to use the broader Heterophasia (including Malacias and Leioptila) and include Mesia in Leiothrix.

Some of the arrangement of species is based on Dong et al. (2010), as is the transfer of several former Actinodura to Ixops (Blyth 1843, type nipalensis).

Turdoidinae Richmond, 1917 (1831)

Leiothrichinae Swainson, 1831

Incertae Sedis

Incertae Sedis: Somewhere in Passerida

1 genus, 1 species

“Incertae sedis” is a fancy term meaning we don't have a clue what to do with these. In this case there is a little clue. Malia is thought to belong to the Passerida somewhere, but its true affinities are currently unknown. It has often been considered a babbler.

Previous Page Next Page