Basal Passeroidea

Core Passeroidea

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

Passeroidea

Passeroidea is the last group of families on the list. It's last mainly because of its size, over 20 families and 1500 species. The first few families involve species that have bounced around the taxonomic tree.

Promeropidae: Sugarbirds Vigors, 1825

1 genus, 2 species HBW-13

The evidence indicates these two African species are basal Passeroidea, although there is a slight chance they outside Passeroidea. Some analyses suggest they are sister to the Modulatricidae (Barker et al., 2004; Beresford et al., 2005; Fig. 3 of Johansson et al. 2008b). Others (Fjeldså et al., 2010; Fuchs et al., 2009; Fig. 2 of Johansson et al. 2008b; Nguembock et al., 2008b) find that Promerops is a separate and deeper independent branch in Passeroidea. Fjeldså et al. (2010) suggested the split might have been 45 million years ago. I think that's somewhat inflated (I don't believe the calibration), but however one dates it, it is an ancient branch. Since it is questionable whether the Promeropidae and Modulatricidae are sisters, and since the division between them is deep even if they are sisters, we treat them as separate families.

Modulatricidae: Spot-throat & allies Fjeldså et al. 2015

3 genera, 3 species Not HBW family

Modulatricidae tree These three birds from sub-saharan Africa have been considered babblers, but they aren't. In fact, they've been bounced around the taxonomic tree. Other suggestions have been that the Kakamega is a thrush, that the Spot-throat is a bulbul or thrush, and that the Dapple-throat is a bulbul or chat. They are none of above and belong in Passerida, next to the sugarbirds (e.g. Johansson et al. 2008b).

Dicaeidae: Flowerpeckers Bonaparte, 1853

3 genera, 49 species HBW-13

Dicaeidae tree The flowerpeckers and sunbirds are sister families (Barker et al., 2004; Ericson and Johansson, 2003, Fjeldså et al, 2010). Some merge them as Nectariniidae (e.g., Sibley and Monroe, 1990; Christidis and Boles, 2008), but Fjeldså et al. found a deep separation between the two clades, deeper than that between Irenidae and Chloropseidae. Nyári et al. (2009) sampled nearly two-thirds of the flowerpeckers. They found two major clades, with members of Dicaeum in both clades. I've moved the Dicaeum in the Prionochilus clade to Pachyglossa (Blyth 1843, type melanoxantha), along with several untested species that are likely also in that group (vincens, annae, aeruginosa, propria).

It's not entirely clear where the Olive-backed Flowerpecker, Prionochilus olivaceus, goes. The Bayesian and maximum likelihood trees in Nyári et al. put it sister to Pachyglossa and the remaining Prionochilus, respectively. That it is basal in the clade cannot be ruled out either. I'm treating it as the basal member of Prionochilus, but it may need its own genus.

Within Dicaeum proper, the anthonyi-quadricolor group is the basal group. The next branch is likely aureolimbatum through retrocinctum, followed by the sister species nigrilore and hypoleucum.

The remaining Dicaeum form a well-supported clade which appears to consist of three subclades: pygmaeum-virescens, sanguinolentum-maugei, and celebicum-tristrami. Note that the Mistletoebird does not appear to be that closely related to the sanguinolentum, even though they have sometimes been regarded as conspecific.

Within the celebicum-tristrami clade, the sister species celebicum and kuehni are basal, followed by another pair, ignipectus and monticolum. Interestingly, all four of these have also been considered conspecific with the Mistletoebird (hirundinaceum). Nonetheless, they also don't seem to be so closely related. Rather, the Mistletoebird seems to be more closely related to the remaining species, and particularly the Red-capped Flowerpecker (geelvinkianum). Only five of the eleven species in the hirundinaceum-tristrami group were sampled, and except for the Mistletoebird, I've adopted the arrangement in HBW, which is consistent with the maximum likelihood tree of Nyári et al. (2009).

The Wakatobi Flowerpecker, Dicaeum kuehni, has been split from Gray-sided Flowerpecker, Dicaeum celebicum, based on Kelly et al. (2014). Splitting allopatric taxa is always tricky as interbreeding potential can often be only indirectly assessed. The two differ in plumage, and mitochondrial DNA differs by over 2.5% (2.53-2.83% for COI). The DNA separation is in a range that offen indicates species status. Moreover, there are noticable plumage and morphological differences as documented by Kelly et al., (2014). To me, this combination justifies the split.

Interestingly, only about 25 miles of open water separates Sulawesi from the Waktobi Islands (part of the Tukangbesi archipelago). The two have never been connected by a ice age land bridge as the water between is a mile or more deep (Google Earth). Kelly et al. found no evidence of recent gene flow in the mitochondrial DNA (COI and ND3). We don't know if any males cross the water and interbred, but it appears no females have recently done so.

Note that I do not make this split based on Hebert et al.'s (2004) barcoding threshold of 2.7%, which I find totally unconvincing. I don't think the authors then had a good understanding of the two separate questions involved: use of barcoding to identify taxa, and use of barcoding to determine species status. Certaintly the sampling and primitive statistical methods used were inadequate to the species status task. This becomes clear from more recent papers such as Tavares and Baker (2008), Kerr et al. (2009), Milá et al. (2012).

Tavares and Baker (2008), who sampled more relevant taxa, found an overlap zone of about 0.8-3.0%. Kerr et al. (2009) studied a large sample of palearctic birds. They looked at type I (false positives) and type II errors (false negatives) from using the barcoding data. They claim an optimum value of 1.6%, although it is rather unclear why this is optimal. Eyeballing their graph (Figure 2) suggests that the minimum total error is at 1.4%. I did not compute the minimum RMS error, but it probably occurs little lower as the RMS criterion puts more weight on equalizing the errors. The errors seem to be equalized at about 1.1%. Taxonomic committees seem to prefer to avoid type I error, with less emphasis on type II error, implying some sort of asymmetric loss function. The optimum error in such cases could be well to right as the type I error region on Kerr et al.'s Figure 2 extends all the way to 8%. The species limits may be incorrect in some of such cases, or the genetic integrity of the species may be maintained through males making the mitochondrial estimates irrelevant (think cuckoos! Fossøy et al., 2010).

Finally, genetic distances in the Neotropics are higher (Milá et al., 2012). To what extent this indicates more actual interspecific variation, or is an artifact of undersplitting remains unclear. I suspect that all this literature could benefit from a more sophisticated approach to both sampling and statistical analysis.

Nectariniidae: Sunbirds Vigors, 1825

16 genera, 143 species HBW-13

Nectariniidae tree The ordering of the sunbirds is based on the tree in Jønsson and Fjeldså (2006a), which is based on Bowie's unpublished Ph.D. dissertation (2003). I've made two alterations, placing Chalcoparia closer to Anthreptes and Leptocoma (see Moyle et. al, 2011), and putting Cinnyris in a position that is a compromise between the Bowie/Jønsson and Fjeldså tree and Nyári et al. (2009b). Checke and Mann (2008) in HBW-13 was used to place the remaining taxa on the resulting tree. There are two exceptions: Loten's Sunbird and the similar Purple Sunbird don't have clear relatives and I doubt they belong in Cinnyris, so I've moved them to the genus Arachnechthra (Cabanis 1851, type lotenia). You'll see it floating at the bottom of the tree.

The basal position of the spiderhunters is clear not only from Bowie/Jønsson and Fjeldså tree and from Nyári et al. (2009b), but also from the the analysis by Moyle et al. (2011), which focuses on spiderhunters. All three sources also also have an oriental clade containing Aethopyga sister to the remaining spiderhunters.

Aethopyga has been rearranged based on Hosner et al. (2013a) and five additional endemic species from the Philippines are recognized:

  1. Crimson Sunbird, Aethopyga siparaja, is split from Magnificent Sunbird, Aethopyga magnifica.
  2. Maroon-naped Sunbird, Aethopyga guimarasensis (including daphoenonota), is split from Flaming Sunbird, Aethopyga flagrans.
  3. Bohol Sunbird, Aethopyga decorosa, and Luzon Sunbird, Aethopyga jefferyi, are split from Metallic-winged Sunbird, Aethopyga pulcherrima.
  4. Tboli Sunbird, Aethopyga tibolii, is split from Apo Sunbird, Aethopyga boltoni.

All of the new species are both genetically and visually distinct. Four have previously been considered separate species. The recently discovered A. tibolii was described as a subspecies (Kennedy et al., 1997), but it too is both genetically and visually distinctive.

Hosner et al. (2013a) also raise the issue of whether the Elegant Sunbird, Aethopyga duyvenbodei, should be placed in a separate genus (Duyvena). Their work weakly supports it as sister to the other Aethopyga, and I leave it in Aethopyga for now.

The reorganization meant subsuming Dreptes into Cinnyris and Drepanorhynchus into Cyanomitra. It also meant resolving any priority issues between Anthodiaeta and Hedydipna by recognizing both. [See Alan Peterson's analysis of the priority issue at zoonomen.net.] The four Asian and Australasian species formerly considered part of Cinnyris have been separated in their own genus, Cyrtostomus.

The genus Cinnyris is now much reduced, as it has additionally lost 15 species to Chalcomitra, 15 to Anthobaphes, and one each to Arachnechthra, Cyanomitra, and Deleornis, gaining only one species from Chalcomitra. The African members of Anthreptes have been dispersed to Deleornis (3) and Hedydipna (7). Nectarinia lost 4 species to Chalcomitra and gains two from Cyanomitra.

Orange-tufted Spiderhunter, Arachnothera flammifera (presumed to include randi), and Pale Spiderhunter, Arachnothera dilutior, have been are split from Little Spiderhunter, Arachnothera longirostra based on the analyses of Lohman et al. (2010) and Rahman et al. (2010). Between them, they consider a decent sampling of the other Little Spiderhunter subspecies, which show relatively little genetic differentiation. This is also supported by Moyle et al. (2011).

Moyle et al. (2011) also supports the split of Bornean Spiderhunter, Arachnothera everetti, from Streaky-breasted Spiderhunter, Arachnothera affinis. Their evidence also suggests that Hypogramma should be subsumed in Arachnothera, as is done here.

Irenidae: Fairy Bluebirds Jerdon, 1863

1 genus, 2 species HBW-10

Whether the fairy bluebirds and leafbirds of the Oriental Region should be united in the same family remains an open question. Most analyses have found they are sister families (Barker et al., 2004; Beresford et al., 2005; Fjeldså et al., 2010; Kennedy et al., 2012; Reddy and Cracraft, 2007; Moltesen et al., 2012); Treplin et al., 2008), although Barker et al. (2002) found differently and Johanssen et al. (2008b) left the issue unresolved. The recent paper by Moltesen et al. suggests that they share a common ancestor a little over 10 million years ago, making the case for treating the fairy bluebirds as a subfamily of the leafbirds. Kennedy et al. (2012) estimated that their common ancestor was likely about 20 million years ago, while Fjeldså et al. put the ancestor around 40 million years ago. In the latter case, they definitely should be separate families.

The Fjeldså et al. estimate is being driven by a single calibration point, a hypothesized origin of Acanthisitta around 82-85 million years ago. I have deep skepticism about this date, and would not be surprised if the true date were much more recent. However, I suspect Moltesen et al. gives too short a time. It seems best to leave these as separate families for now.

I'm assuming that the flowerpecker/sunbird clade is a slightly deeper branch than the fairy bluebird/leafbird clade. However, although it seems to be the consensus of the relevant papers, the overall support for this is not strong.

Moltesen et al. (2012) advocate the treatment of a number of Irena and Chloropsis subspecies as full species, primarly on the basis of genetic distance. I think they are likely right about some of them, but prefer to wait for more evidence in most cases.

Chloropseidae: Leafbirds Wetmore, 1960 (1847)

1 genus, 12 species HBW-10

I find one of the Moltesen et al. (2012) splits more compelling as they also found that one of the subspecies was more closely related to an entirely different species. As a result, I've separated the Yellow-bordered Leafbird, Chloropsis septentrionalis from the Lesser Green-Leafbird, Chloropsis cyanopogon. There didn't seem to be a historical English name for septentrionalis, so I've given it one based on a distinguishing characteristic, the yellow border around the mask on the males, as noted by Moltesen et al.

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