Winter Wren potential split (1 Viewer)

Hi

This link was recently posted on Frontiers of Bird ID.

http://www.blackwell-synergy.com/doi/abs/10.1111/j.1365-294X.2008.03769.x

Maybe the Scottish Wrens will be up for detailed study next. We could do with an endemic species in Scotland...

See also Drovetski et al (2004), which identified 6 clades: W Nearctic, E Nearctic, E Asia, Nepal, Caucasus, Europe (presumably pacificus, hiemalis, fumigatus, nipalensis, hyrcanus, troglodytes respectively).

http://www.pubmedcentral.nih.gov/picrender.fcgi?artid=1691619&blobtype=pdf

Richard

A look at Scottish wrens may well be on the cards, there was some interest a couple of years ago but the project didn't get off the ground (lack of funding possibly) - the theory was that St Kilda and Fair Isle wrens may well be candidates for species.

Cheers,
Andrew

...and the distinctive Pribilof (alascensis) and Aleutian (meligerus) forms surely also warrant further study (both earlier recognised as species by AOU until lumped with Winter Wren in the 1931 checklist).

Richard

Richard Klim said:

...and the distinctive Pribilof (alascensis) and Aleutian (meligerus) forms surely also warrant further study (both earlier recognised as species by AOU until lumped with Winter Wren in the 1931 checklist).

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I looked up the electronic appendix to Drovetski et al (2004)... both alascensis and meligerus were sampled in the phylogeny, and are in the western nearctic clade, which shows essentially no variation throughout. It's interesting how none of the six major clades shows any significant variation, actually. As far as the Scottish wrens go, there are six samples identified as "GB" from indigenus which I assume is Great Britain, but they're not identified to locale beyond that.

~ Nick

ntbirdman said:

... both alascensis and meligerus were sampled in the phylogeny, and are in the western nearctic clade, which shows essentially no variation throughout. It's interesting how none of the six major clades shows any significant variation, actually.

~ Nick

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Oh well...

But while watching an alascensis Wren on St. Paul Island last June, it seemed incredible that the large and sluggish bird creeping furtively through the grass at my feet (rather like a Lanceolated Warbler) could really the be the same species as those that dash energetically around my garden back home in England.

Richard

ntbirdman said:

I looked up the electronic appendix to Drovetski et al (2004)... both alascensis and meligerus were sampled in the phylogeny, and are in the western nearctic clade, which shows essentially no variation throughout. It's interesting how none of the six major clades shows any significant variation, actually.

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Drovetski et al. had three samples from this area, one alascensis from St Paul ('sp1'), one meligerus from Attu ('rwd24877') and one tanagensis from Adak ('ksw2720'). These are very close to the rest of the pacificus clade but, interestingly, they group together in the phylogenetic tree shown in the paper.
See also http://dx.doi.org/10.1007/s10584-007-9332-6 (pdf currently freely available) - the birds of Attu, Adak and Commander Is (pallescens) belong to the W nearctic group (which, as a passing note, is therefore not strictly nearctic), but form a distinct cluster within this group and do not seem to share any haplotype with the birds of mainland AK and Kodiak.
The distance is too small to suggest species status, but this still hints at some level of isolation/differentiation.

Data from Kamchatka would be interesting too. The birds there are said by Dementiev to be the same race as those of Commander - if true, and if the latter belong to the W nearctic clade, it might be that there is an actual contact zone between this clade and the Asian clade to be found somewhere up there...

As far as the Scottish wrens go, there are six samples identified as "GB" from indigenus which I assume is Great Britain, but they're not identified to locale beyond that.

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indigenus is the subspecies found on mainland GB and Ireland, and is rather poorly differentiated from nominate troglodytes (particularly in the southern part of its range). I don't think there are any published genetic data pertaining to the northern island populations, among which are the morphologically more distinct taxa.

islandicus from Iceland would surely deserves attention, too.

Cheers,
Laurent -

I have only gotten through the 2004 paper so far, but one interesting little note in their text was that the closests relative of Winter Wren was not House Wren but Marsh and Sedge Wrens. Any opinions about what that eventually will lead to regarding which genus these species belong to?

On a related note, is there any new information out there for House Wren?

Edit: one particular interest for me, as I live in the Caribbean is the island populations that is currently placed in House Wren. Any thought on these?

Thanks
Niels

I have used Nannus troglodytes as the scientific name for years, but I'm not sure how many others do this.

The Marsh and House Wrens can then stay where they are!

Frenchy said:

Hi

This link was recently posted on Frontiers of Bird ID.

http://www.blackwell-synergy.com/doi/abs/10.1111/j.1365-294X.2008.03769.x

Maybe the Scottish Wrens will be up for detailed study next. We could do with an endemic species in Scotland...

Click to expand...

Anyone got a full Pdf of this paper - I am too tight to pay for it !

Lindsay

njlarsen said:

I have only gotten through the 2004 paper so far, but one interesting little note in their text was that the closests relative of Winter Wren was not House Wren but Marsh and Sedge Wrens. Any opinions about what that eventually will lead to regarding which genus these species belong to?

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The first file in attachment is a ML tree based on ND2 sequences from GenBank. I've tried several other genes (cytochrome b, COI, beta-fibrinogen intron 4) and, as far as generic limits would be concerned, they give similar results - but ND2 has a broader taxonomic sampling, hence is the most interesting to show.

Troglodytes s.l., Thryorchilus and Cistothorus form a clade that has a very strong support. This clade includes three main, well-supported lineages, namely [Troglodytes s.s. + Thryorchilus], Cistothorus, and Nannus.
The basal node in this clade groups Cistothorus with Nannus in most cases, but always with very poor support; in some trees I get a node grouping Cistothorus with [Troglodytes + Thryorchilus] instead (see also http://www.specifysoftware.org/Informatics/bios/biostownpeterson/MBP_A_2005.pdf).
Thryorchilus may be basal to all Troglodytes s.s., but the node grouping the latter is also poorly supported.

Based on this, several options could be possible, but the most straightforward would indeed probably be to recognize the three main lineages, because they have strong individual support, thus transfer the winter wren to Nannus. This move would be rather secure if it can gain wide acceptance (the probability that the resulting taxonomy could be proved unacceptable would be very low) - the question might be whether the (huge) part of the World that has a single wren species to deal with, would be ready to accept it.

Few people use Nannus currently, I'm afraid - just compare the number of hits in Google -
http://www.google.be/search?q=wren+nannus-troglodytes
http://www.google.be/search?q=wren+troglodytes-troglodytes

On a related note, is there any new information out there for House Wren?

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Note the structure within the house wren group in the ND2 tree: brunneicollis is basal to the house wren complex; two sequences of aedon from Illinois group with a musculus sequence (from Oaxaca, Mexico); one sequence of aedon from Washington State groups with the Socorro wren.

The second attached file is a COI tree generated by feeding the BOLD ID engine (http://www.barcodinglife.org) with a house wren COI sequence from GenBank. I used a full-database search, which means that the tree can include unvalidated sequences, some of which may be misidentified, hence we have to look at it with caution. (In particular, if a taxon pops up in a tree at a completely unexpected place, this is certainly not to be trusted.)

It is rather clear that there is a lot of genetic structure in the house wren (which parallels the case of the winter wren).

The most divergent bird labelled T. aedon in the BOLD database is from Venezuela. It differs from the focal sequence I used to generate the tree by 8.3% (for comparison, the largest distances within T. troglodytes barely pass 6%). It clusters with (but is still quite distant from) sequences of T. solstitialis from Argentina: this is quite odd for a house wren, so I'm not sure we should buy this one. (This could conceivably be a misidentified T. solstitialis from another population, or a T. rufulus, both of which occur in Venezuela.)

The next most divergent bird is still 7.4% away from the focal sequence. This one clusters with a sequence of Henicorhina in the tree, which is clearly odd too - but in this case, the Henicorhina ID is probably the first one we should question, as this genus should not normally appear in this tree at all, certainly not in a position embedded in Troglodytes. Both samples are from SC Mexico. Brunneicollis? The phylogenetic position would in any case be consistent with that of this taxon in the ND2 tree, and the location would seem to fit as well.

Then we get a group of three clusters, two of which correspond to populations usually treated as different species, T. sissonii (Socorro wren) and T. tanneri (Clarion wren), and the last being composed of T. aedon from W North America (British Columbia), together with a bird from C Mexico. The divergence from the focal sequence there is around 5.5-6%, comparable to the divergence between the W and E American groups of T. troglodytes.

The remaining includes, in order of decreasing apparent divergence from the focal sequence:
- a group of two clusters: one from the Yucatan/Quintana Roo in SE Mexico; the other from Dominica;
- a cluster from S Argentina (Tierra del Fuego, Sta Cruz, Rio Negro);
- a cluster from C and N Argentina (Buenos Aires, Corrientes, Formosa, Entre Rios, Jujuy);
- a group of four clusters: one from N Argentina (Formosa, Misiones); one from Grenada; and two from Trinidad & Tobago, one of which also includes a bird from Venezuela; and
- a cluster from NC/NE America (Quebec, Ontario).
Divergence levels withing this entire group are below 5%.

The usual cautionary statements apply - this tree is based on data, and generated with a method, that are aimed at identifying clusters of closely related haplotypes, not relationships. No measures of support are given, and (some of the) the relationships shown in the tree can be wrong. But I just can't go without noticing that the grouping of British Columbia aedon with T. sissonii parallels the grouping of a Washington aedon with the same taxon in the ND2 tree; and the grouping of Ontario aedon with birds of Yucatan parallels the grouping of two Illinois aedon with a T. (aedon) musculus from Oaxaca in the ND2 tree.
This means that we have two independent data sets that suggest congruently that the deepest genetic divergence in the house wren complex, if you exclude brunneicollis, could be between two North American populations....

Laurent,

This is great. I cannot open the first attached file (ND2 tree) though.

For the House Wren, there is also this A.O.U. 124th meeting abstract (do not know whether a paper has been published or is in progress) held on October 3-7, 2006 in Veracruz, Mexico:

Klicka, J., Barrick Museum, University of Nevada - Las Vegas, Las Vegas, NV, USA, [email protected];
Bermingham, E., Smithsonian Tropical Research Institute, Balboa, Panama, Panama;
Ricklefs, R. E., Department of Biology, University of Missouri - St. Louis, St. Louis, MO, USA;
Escalante-Pliego, P., Instituto de Bilogia, Universidad Nacional Autonoma de Mexico, DF, Mexico;
Spellman, G. M., Barrick Museum, University of Nevada - Las Vegas, Las Vegas, NV, USA

A SYSTEMATIC ASSESSMENT OF THE HOUSE WREN SPECIES COMPLEX

The House Wren (Troglodytes aedon) has the largest breeding distribution of any New World songbird, occurring from Southern Canada to Tierra del Fuego. Although currently recognized as a single species it is widely regarded as a complex comprised of several historically recognized forms, many of which have previously been considered species. These include the nominate “race”, aedon (Northern House Wren; North America, north of Mexico), brunneicollis (Brown-throated House Wren ; Mexican highlands), and musculus the (Southern House Wren; Southern Mexico through South America). The complex also includes a number of morphologically distinctive island populations including martinicensis (Lesser Antilles), beani (Isla Cozumel), and tanneri (Isla Clarion). Previous molecular studies on this assemblage have led to conflicting results and a comprehensive analysis including representation from all of these groups is lacking. Using mtDNA sequence data (complete ND2 and ATPase[6&8] genes), dense taxon sampling (n = 188), and modern analytical methods, we reconstruct a phylogentic hypothesis for members of the House Wren complex. Prelminary analyses indicate that the Northern House Wren (aedon) as presently recognized, is polyphyletic. Western house wrens (T. aedon parkmanii) form a well-supported clade that is sister to the Clarion Island form, tanneri. Eastern house wrens (T. aedon aedon) are part of a “low-land” house wren radiation that includes the southern (musculus), Lesser Antillean (martinicensis) and Cozumel (beani) forms. Our data suggest multiple colonizations of the lesser Antilles from northern South America. Although sampling is limited, populations on Dominica (n = 4), St. Vincent (n = 3), and Grenada (n = 4) appear to be reciprocally monophyletic whereas Trinidad (n = 12) shares haplotypes with the South American mainland. The Mexican highland form brunneicollis diverged early in the history of the complex and is sister to all other groups. It too is polyphyletic with individuals from Sierra Madre Occidental lying outside of an otherwise well-defined clade.

Hello Daniel,
Somehow I had overlooked this abstract, obviously, thanks for this. Interesting - and largely congruent with the rest. So it makes three data sets, now

I cannot open the first attached file (ND2 tree) though.

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I've replaced it with another one (in my original message, above). TreeFinder produces PostScript files, that any basic Linux/Unix machine should be able to read, but many Windows machines can't. It seems my file was still a bit "too much PostScript" for Acrobat... Does this one work?

L -

Thank you Laurent and Daniel,
a lot of work that my little question generated! I am truly looking forward to study this a little more when I have some time, hopefully this weekend.

Niels

BNA Online

Toews & Irwin 2012. Pacific Wren (Troglodytes pacificus). BNA Online.

Geographic Variation

Phenotypic variation in the species is complex and involves differences in size and proportions—notably wing chord, tail length, body mass, and relative size of bill, tarsus, and toes—and plumage coloration and pattern, particularly in the depth and hue of the throat and dorsum, as well as the extent and width of barring and spotting on the back. In general, birds in Alaska are largest, particularly those on the w. Aleutians and the Pribilofs. Birds on the mainland of the Pacific Northwest and in w. Canada are the most rufescent, with populations becoming progressively browner then grayer to the west. In addition to being either grayer or browner, populations in Alaska also tend to be paler.

Phenotypic differentiation apparently has been recent and rapid given that there is little differentiation in neutral markers, in this case cytochrome b from the mitochondrial DNA (mtDNA), from Kodiak through the Aleutian chain to the Commander Is. (Pruett and Winker 2008). Likewise, populations of two subspecies in se. British Columbia have yet to diverge at neutral sites in the mtDNA or nuclear genome (Toews and Irwin 2008).

Subspecies

Seven subspecies, following Phillips (1986), as amended by Browning (1990). Many of the described subspecies are from insular Alaska, and the diagnosability of some taxa remains an open question. It is clear that "All Alaska material from east of the Aleutians needs to be assembled in one place and reviewed" (Gibson and Kessel 1997). As a case in point, although Browning (1990) suggested three putative subspecies described by Amadeo M. Rea (in Phillips 1986) are likely to be valid, he cautioned that their validity "cannot be confirmed until a thorough analysis of the geographic variation of the western populations is presented" and recommended that until such an analysis appears these taxa be treated as junior synonyms of the nominate subspecies. ...

[pacificus, salebrosus, helleri, alascensis, semidiensis, kiskensis, meligerus.]

Related Species

... Whether the genus Troglodytes, as currently constituted, is monophyletic is open to question, but some (e.g., Rice et al. 1999) have suggested that the T. troglodytes complex—which includes the nominate group in Eurasia as well as T. pacificus and T. hiemalis, the Winter Wren of e. North America—ought to be placed in a separate genus, as it once was (e.g., Ridgway 1904).

This species complex was considered to be a single species for many decades, lumped under the scientific name T. troglodytes and English names of the Winter Wren or, in Europe, simply the Wren. Yet patterns of genetic and vocal variation in an area of overlap between western and eastern populations in North America implied that the populations are reproductively isolated (Toews and Irwin 2008, Chesser et al. 2010). Populations across Eurasia, which are allopatric from North American populations, also were considered to be specifically distinct because a phylogeny based on mtDNA placed them nearer to populations in e. North America than to those in w. North America (Drovetski et al. 2004, Chesser et al. 2010). Divergence dates, on the basis of mtDNA sequence changes, estimate that T. pacificus split from the other two taxa 4 mya (Drovetski et al. 2004, Toews and Irwin 2008), and that T. hiemalis and T. troglodytes sensu stricto diverged from each other 2 mya, a curious pattern in given that T. pacificus and T. hiemalis occur on the same continent.

Morphological intermediates involving T. pacificus are unknown, although its phenotype is near enough to that of T. hiemalis that intermediacy would be difficult to detect. Moreover, Toews and Irwin (2008) reported that a male collected near Gavin Lake, British Columbia, carried mtDNA of T. hiemalis but had a nuclear genome intermediate between T. hiemalis and T. pacificus, suggesting it was a first generation hybrid. Still, genetic patterns between the two species indicate that interbreeding is rare and does not result in noticeable gene flow (Toews and Irwin 2008).

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Toews & Irwin 2008.

Shannon T.J., McGowan R.Y., Zonfrillo B., Piertney S., Collinson J.M. (in press): A genetic screen of the island races of Wren Troglodytes troglodytes in the North-east Atlantic. Bird Study. [abstract]

Capsule Atlantic Island Wrens are very closely related to mainland European populations.
Aims The first genetic screen of the North-east Atlantic island subspecies of (Winter) Wren Troglodytes troglodytes was performed to resolve their relationship to mainland Eurasian and Nearctic populations.
Methods The ND2 gene was sequenced from 15 wrens from Iceland, Faroe Islands, St Kilda, Outer Hebrides, Fair Isle and Shetland (2–3 individuals of each subspecies) and compared with Eurasian Wrens from mainland Britain and Europe, and Winter Wrens from North America.
Results All island subspecies were shown to originate from European rather than Nearctic founders. Genetic divergence from mainland British and European populations was small (0.1–0.3% uncorrected). The major European haplotype was present in some individuals from Shetland, Fair Isle and Faroes. Novel unique haplotypes were found in all individuals of St Kilda, Iceland and Hebridean Wrens, and in two individuals of Fair Isle Wren, contrasting with the high inferred levels of gene flow across Europe.
Conclusions Genetic data are consistent with a postglacial colonization of Atlantic islands from mainland Britain and Europe, possibly with continued gene flow due to migration of European birds. Tentatively, St Kilda Wren and Iceland Wren may be more closely related; most other subspecies are probably polyphyletic.

Japan

Asai, Shigeta, Saitoh & Iwami. Phylogeny and validity of Eurasian Wren Troglodytes troglodytes subspecies breeding in Japan. 26th IOC, Tokyo, 2014.

The Check-List of Japanese Birds recognizes three subspecies of Eurasian Wren Troglodytes troglodytes that breed in Japan [= fumigatus, mosukei, ogawae], and one extinct subspecies [= orii]. In the Check-List, the subspecies breed on the mainland, including Hokkaido and Kyushu, on the southern islands of the Izu Islands, and in the Osumi Islands, respectively, whereas Momiyama (1927) subdivided them and recognized four additional subspecies that breed in the following regions: Hokkaido, western Honshu, northern Kyushu, and southern Kyushu. According to the original descriptions of the subspecies, they were proposed from one to three samples that were captured in January, February, March, or December, which is considered to be the nonbreeding season. Therefore, their localities may not be appropriate if the individuals had migrated. Furthermore, given that their descriptions depended on plumage color, the sample sizes may have been too small. The latter four subspecies are thought to be synonyms of T. t. fumigatus because of these concerns regarding reliability. Nevertheless, no one has sufficiently examined the validity of those subspecies. In this study, we examined whether the putative subspecies from the sampling sites are monophyletic using molecular phylogenetic methods. We used blood or muscle samples collected from May to August, during the tentative breeding season. Complete cytochrome b and control regions in mitochondrial DNA were sequenced and the best substitution models were selected. Phylogenetic trees were constructed using Maximum Likelihood (ML) and Bayesian methods with Eurasian Treecreeper Certhia familiaris sequences as an outgroup. Although bootstrap values for the ML tree were low overall, both trees showed that three samples collected from the southern Izu Islands were monophyletic. Two samples collected from Yaku Island, of the Osumi Islands, were paraphyletic in the Bayesian tree but not in the ML tree. Therefore, T. t. mosukei breeding in the Izu Islands is most likely valid. T. t. ogawae needs to be re-examined with more samples.

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Kroodsma & Brewer 2005 (HBW 10).

[See also: Daito Wren.]

Xenospiza said:

I have used Nannus troglodytes as the scientific name for years, but I'm not sure how many others do this.

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l_raty said:

Troglodytes s.l., Thryorchilus and Cistothorus form a clade that has a very strong support. This clade includes three main, well-supported lineages, namely [Troglodytes s.s. + Thryorchilus], Cistothorus, and Nannus.
The basal node in this clade groups Cistothorus with Nannus in most cases, but always with very poor support; in some trees I get a node grouping Cistothorus with [Troglodytes + Thryorchilus] instead (see also http://www.specifysoftware.org/Informatics/bios/biostownpeterson/MBP_A_2005.pdf).
Thryorchilus may be basal to all Troglodytes s.s., but the node grouping the latter is also poorly supported.

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Why Nannus? After all, the type species of Troglodytes is, by tautonymy, Troglodytes troglodytes. So if the genus is to be split, it should be T. aedon and relatives that are moved to a new genus, not T. troglodytes.

Similar to the erroneous transfer by Madge & Burn (Wildfowl) of Whooper, Trumpeter and Tundra Swans to Olor, despite Whooper Cygnus cygnus being the type of Cygnus.

Nutcracker said:

Why Nannus? After all, the type species of Troglodytes is, by tautonymy, Troglodytes troglodytes. So if the genus is to be split, it should be T. aedon and relatives that are moved to a new genus, not T. troglodytes.

Similar to the erroneous transfer by Madge & Burn (Wildfowl) of Whooper, Trumpeter and Tundra Swans to Olor, despite Whooper Cygnus cygnus being the type of Cygnus.

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What are the sources you are using to make these statements?

I think that you'll find that the 'double name' does not automatically mean that this species is the type for the genus of the same name. Often the situation with very old (thus, mainly European) species descriptions involved those species being 'forced' into a very limited group of genera originally described by Linnaeus and Gmelin, and only placed in separate genera several decades later, as the field of taxonomy grew and our understanding of relationships improved. Originally, according to Peters and Hellmayr, T. troglodytes was described as Motacilla troglodytes (Linnaeus 1758) and C. cygnus was described as Anas cygnus (Linnaeus 1758). T. aedon is the type species of the genus Troglodytes Vieillot 1808, whereas Nannus Billberg 1828 was coined for T. troglodytes. In the cases of the swans, Cygnus Bechstein 1803 was first coined for C. olor, whereas Olor was created for C. cygnus by Wagler 1832.

Cygnus Bechstein, 1803 [OD].
Originally included species:

• Anas Olor Gmelin [OD]

Anas cygnus Linnaeus is not cited, thus it can definitely not be the type by tautonymy. Anas Olor Gmelin is the type species by original monotypy.
(Note that there are also suggestions that Cygnus might be regarded as available from Garsault 1764 ["OD"]. This would not change anything regarding the type species, though.)


Olor Wagler, 1832 [OD].
Originally included species:

• Olor musicus (Cygnus musicus Bechstein)
• Olor Bewickii (Cygnus bewickii Yarrell)
• Olor Buccinator (Cygnus Buccinator Richardson)

Anas olor Gmelin is not cited, thus it can definitely not be the type by tautonymy. (And the descriptive text [die Stirn ohne höcker: the forehead without a bump] actually excludes it.) No type species fixed in the OD. Type species deemed (eg., Richmond, AOU 1998) to have been fixed by subsequent designation by Gray 1840 as Cygnus musicus Bechstein [OD], which is a junior synonym of Anas cygnus Linnaeus. Gray did not do this, though: he designated "Cygnus mansuetus" (L.) Wagler", adding a reference to a plate showing a Mute Swan (Planche enluminée 913). "Cygnus mansuetus" is a prelinnean name, used by Willughby and Ray for the Mute Swan, and cited by Linnaeus 1758 under the varietas β of his Anas cygnus [here]. This designation is invalid (what Gray designated was not an originally included species); I'm unsure who actually designated a type validly.


Troglodytes Vieillot, 1809 [OD].
Originally included species:

• Troglodytes aedon Vieillot
• Troglodytes arundinaceus Vieillot

This case is less straightforward than the previous two, because Motacilla troglodytes Linnaeus/Gmelin is cited in the synonymy of Troglodytes arundinaceus Vieillot [here], and this might be interpreted as satisfying Art.68.4: "If a valid species-group name, or its cited synonym, originally included [Art. 67.2] in a nominal genus-group taxon is identical with the name of that taxon, the nominal species denoted by that specific name (if available) is the type species (type species by absolute tautonymy)", effectively making Motacilla troglodytes Linnaeus the type species by absolute tautonymy. What Troglodytes arundinaceus is deemed a synonym of, however, is the "Var. Y" (actually the varietas γ) of M. troglogytes, described but not named by Gmelin 1789 [here]; Motacilla troglodytes itself is not presented as "a cited synonym" of "a valid species-group name" originally included in the genus, and therefore does not qualify as the type species.
No type species fixed in the OD. Type species fixed as Troglodytes aedon Vieillot by subsequent designation by Baird 1858.