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Calandrella

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Old Saturday 16th August 2014, 08:26   #1
Daniel Philippe
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Calandrella

Redrawing the phylogeny and taxonomy of Calandrella Larks: one new localized West African taxon and several surprising relationships

M. Stervander, U. Olsson, P. Alström, U. Ottosson, B. Hansson, and S. Bensch

Larks are known as a morphologically diverse family which has long challenged taxonomists. Recently, a multilocus phylogeny of all genera, and >80 % of the species, revealed several surprising relationship warranting a major taxonomical revision. We studied the genus Calandrella (excluding the paraphyletic Alaudala clade sensu Alström et al. 2013), covering all species and subspecies currently recognized by IOC. Sampling wild birds we sequenced a large portion of, or the entire, mitochondrial cytb gene. This was supplemented by sequencing a 350 bp region from museum specimens. The results reveal some highlights: (1) A localized population of Red-capped Lark C. cinerea in Nigeria is a unique, non-recognized taxon, which is sister to the subspecies C. c. williamsi in East Africa. This clade differs substantially from all other C. cinerea populations in Southern Africa, which in turn make up a panmictic population. (2) In the Horn of Africa and Arabian Peninsula, C. erlangeri and C. b. blanfordi are recent sister taxa, as are C. b. daaroodensis and C. b. eremica, whereas these two clades represent deeply split sister lineages. (3) The easternmost subspecies dukhunensis of Short-toed Lark C. brachydactyla groups as sister to the Hume’s Short-toed Lark C. acutirostris. Remaining Short-toed Lark populations roughly differentiate into a Western (brachydactyla) and an Eastern (longipennis) clade, albeit with some mixing. For verification, we performed Restriction site Associated DNA (RAD) sequencing on single representatives for all major clades, yielding some 70,000 orthologous loci across the genome. Phylogenetic modelling of single nucleotide polymorphism data from the RAD dataset was performed to compare to the patterns obtained from the single locus mitochondrial analysis. We will evaluate the use of RAD sequencing for phylogenetic inference, and the possibility to use degraded tissue from museum specimens in RAD sequencing.

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Old Saturday 16th August 2014, 09:56   #2
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Greater Short-toed Lark

Quote:
Originally Posted by Daniel Philippe View Post
We studied the genus Calandrella (excluding the paraphyletic Alaudala clade sensu Alström et al. 2013), covering all species and subspecies currently recognized by IOC. ... (3) The easternmost subspecies dukhunensis of Short-toed Lark C. brachydactyla groups as sister to the Hume’s Short-toed Lark C. acutirostris. Remaining Short-toed Lark populations roughly differentiate into a Western (brachydactyla) and an Eastern (longipennis) clade, albeit with some mixing.
The deep divergence of dukhunensis was discussed in Alström et al 2013, but it would be interesting to know the approximate geographic scope of the longipennis clade. It must at least include orientalis, which is anyway synonymised with longipennis in de Juana & Suárez 2004 (HBW 9).

Incidentally, of the other subspecies recognised by IOC, woltersi is synonymised (with hermonensis?) in Kirwan et al 2008 (Birds of Turkey).
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Old Thursday 26th May 2016, 07:46   #3
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Martin Stervander, Per Alström, Urban Olsson, Ulf Ottosson, Bengt Hansson, Staffan Bensch. Multiple instances of paraphyletic species and cryptic taxa revealed by mitochondrial and nuclear RAD data for Calandrella larks (Aves: Alaudidae). Molecular Phylogenetics and Evolution. In Press, Accepted Manuscript, Available online 26 May 2016.

[abstract]
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Old Thursday 26th May 2016, 08:49   #4
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Quote:
Originally Posted by Peter Kovalik View Post
Martin Stervander, Per Alström, Urban Olsson, Ulf Ottosson, Bengt Hansson, Staffan Bensch. Multiple instances of paraphyletic species and cryptic taxa revealed by mitochondrial and nuclear RAD data for Calandrella larks (Aves: Alaudidae). Molecular Phylogenetics and Evolution. In Press, Accepted Manuscript, Available online 26 May 2016.

[abstract]
New species or subspecies elevated to species rank ?
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Old Thursday 26th May 2016, 09:13   #5
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Originally Posted by Nomenclatorist View Post
New species or subspecies elevated to species rank ?
Stervander et al.:
Quote:
Results:
Clade A comprised all C. brachydactyla subspecies except C. b. dukhunensis.
Clade B comprised the Asian C. acutirostris.
Clade C contained our single sample of C. brachydactyla dukhunensis
Clades D and E contained the sub-Saharan C. cinerea, with a deep split, estimated at 2.7 mya (95% HPD 1.7–3.7 mya) that separated the Nigerian C. c. saturatior (D1) and Kenyan C. c. williamsi (D2) from the South African C. c. cinerea and DR Congo C. c. saturatior (E).
Clade F included C. b. blanfordi (N Eritrea; F1) and C. erlangeri (Ethiopia; F2), separated from clade G, which contained C. blanfordi eremica (Arabia; G1; one sample (44_SAU) inconclusively placed within clade G in ML analyses) and C. b. daaroodensis (N Somalia; G2), 4.3 mya (95% HPD 3.0–5.6 mya).
Quote:
Conclusions
...In the genus Calandrella, we discovered several cryptic lineages within C. acutirostris and C. cinerea, whereas many described subspecies were not genetically distinct; the family is thus in need of taxonomic revision, which we will address elsewhere.
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Old Thursday 26th May 2016, 09:49   #6
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Quote:
Originally Posted by Peter Kovalik View Post
Martin Stervander, Per Alström, Urban Olsson, Ulf Ottosson, Bengt Hansson, Staffan Bensch. Multiple instances of paraphyletic species and cryptic taxa revealed by mitochondrial and nuclear RAD data for Calandrella larks (Aves: Alaudidae). Molecular Phylogenetics and Evolution. In Press, Accepted Manuscript, Available online 26 May 2016.
[Fig. 1]
[Fig. 2]
[Fig. 3]
[Fig. 4]
[Fig. 5]

For the clades referred to in the text quoted in Peter's last post, see Fig. 2 (complete cyt-b sequences) and 3 (complete + shorter cyt-b sequences).

Last edited by l_raty : Thursday 26th May 2016 at 09:57.
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Old Sunday 29th May 2016, 20:31   #7
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Quote:
Originally Posted by Peter Kovalik View Post
Martin Stervander, Per Alström, Urban Olsson, Ulf Ottosson, Bengt Hansson, Staffan Bensch. Multiple instances of paraphyletic species and cryptic taxa revealed by mitochondrial and nuclear RAD data for Calandrella larks (Aves: Alaudidae). Molecular Phylogenetics and Evolution. In Press, Accepted Manuscript, Available online 26 May 2016.

[abstract]
TiF Update May 29, 2016

Calandrella Larks: The genus Calandrella has been studied in detail by Stervander et al. (2016). I use their arrangment, and have implemented three splits.

Darod Lark, Calandrella daaroodensis, of northern Somalia has been split from Blanford's Lark, Calandrella blanfordi.
Red-capped Lark, Calandrella cinerea, is split into Northern Red-capped Lark, Calandrella williamsi, and Southern Red-capped Lark, Calandrella cinerea. The Northern Red-capped Lark only the Kenyan birds (williamsi) and Nigerian birds usually attributed to saturatior. If they had a scientific name, I would probably split the Nigerian and Kenyan birds into two species. The type of saturatior is from Angola, representing a Southern Red-capped Lark. The isolated northern populations in the DR Congo and Uganda belong in Southern Red-capped Lark.
Sykes's Short-toed Lark, Calandrella dukhunensis, has been split from Greater Short-toed Lark, Calandrella brachydactyla. The name Rufous Short-toed Lark has been used for this species, but has also been used more appropriately for the Somali Short-toed Lark, Alaudala somalica.
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Old Saturday 9th July 2016, 06:08   #8
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TiF Update July 8, 2016

Larks: When separating Darod Lark, Calandrella daaroodensis, from Blanford's Lark, Calandrella blanfordi, I had missed that eremica belongs with daaroodensis. Since eremica has priority, Darod Lark becomes Calandrella eremica. Since the resulting species occurs on both sides of the Gulf of Aden, Darod Lark is no longer an appropriate name, and no name seems to be in use, I'm calling it Aden Lark. In sum, Darod Lark, Calandrella daaroodensis is replaced by Aden Lark, Calandrella eremica.
[Alaudidae, Paroidea & Sylvioidea I, 3.08]
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Old Wednesday 24th August 2016, 06:17   #9
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Calandrella dukhunensis

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Originally Posted by Peter Kovalik View Post
Martin Stervander, Per Alström, Urban Olsson, Ulf Ottosson, Bengt Hansson, Staffan Bensch. Multiple instances of paraphyletic species and cryptic taxa revealed by mitochondrial and nuclear RAD data for Calandrella larks (Aves: Alaudidae). Molecular Phylogenetics and Evolution. In Press, Accepted Manuscript, Available online 26 May 2016.

[abstract]
IOC Updates Diary Aug 23
Post proposed split of Syke’s Short-toed Lark on Updates/PS

Calandrella dukhunensis is recognized by TiF v3.06 and The eBird/Clements v2016.
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Old Wednesday 24th August 2016, 09:58   #10
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Quote:
Originally Posted by Peter Kovalik View Post
IOC Updates Diary Aug 23
Post proposed split of Syke’s Short-toed Lark on Updates/PS

Calandrella dukhunensis is recognized by TiF v3.06 and The eBird/Clements v2016.
It should be "Sykes's Short-toed Lark", not '"Syke's" - I notice IOC get it wrong, too - the surname was "Sykes" - however, I prefer my Scottish education's version, "Sykes' Short-toed Lark", because the odd custom in England is to pronounce the possessive of Sykes as "Sykes-zs"!
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Species and subspecies are but a convenient fiction - Kees van Deemter (2010), "In praise of vagueness". Biology is messy
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Old Thursday 22nd September 2016, 19:46   #11
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Hi guys,

Nice to see you immediately picked up our Calandrella paper once it became available. I intended to comment in this thread earlier, but I never got around to do so, and by now most of what I wanted to point out was also figured out by John Boyd (TiF) and discussed here.

I can tell you that we are preparing a manuscript that deals with the morphometrics and the taxonomy of Calandrella, which I hope that we will be able to submit during the autumn. Nevertheless, the general gist of the taxonomic consequences are the ones discussed above (though exact boundaries between species and subspecies rank are not obvious in all cases).

If anyone feels like reading a popular science summary of the published paper, you are welcome to have a look at my site.

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Old Thursday 6th October 2016, 11:40   #12
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Calandrella dukhunensis

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Originally Posted by Peter Kovalik View Post
Martin Stervander, Per Alström, Urban Olsson, Ulf Ottosson, Bengt Hansson, Staffan Bensch. Multiple instances of paraphyletic species and cryptic taxa revealed by mitochondrial and nuclear RAD data for Calandrella larks (Aves: Alaudidae). Molecular Phylogenetics and Evolution. In Press, Accepted Manuscript, Available online 26 May 2016.

[abstract]
IOC Updates Diary Oct 5

Accept Mongolian Short-toed Lark
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Old Friday 23rd June 2017, 07:53   #13
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"Tephrocorys ruficeps" in Shelly 1902 ... ?

Anyone know which of today's taxon is equal of the "T. ruficeps" [Tephrocorys ruficeps], mentioned in the OD of Calandrella blanfordi SHELLEY 1902 (here, & plate, here, fig.2) as "Tephrocorys blanfordi" (where Dr. Blanford was the first to regard the latter one "as distinct from T. ruficeps") ... ?

I assume that this "ruficeps" today is equal of a Calandrella, but which one?

I can´t find it on either level in the IOC list, ver. 7.1 [that the Swedish World list today (mostly) follows].

Is it an invalid name for today's Erlanger's Lark Calandrella (blanfordi) erlangeri NEUMANN 1906 (earlier on a subspecies of Red-capped Lark C. cinerea GMELIN 1789), or ... ?

Anyone know?

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Old Friday 23rd June 2017, 08:17   #14
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Tephrocorys ruficeps.
According to Sharpe 1890, Cat. XIII, p. 563, Alauda ruficeps Rüppell, 1835, inhabits "N. E. Africa"; from the description this seems to be a synonym for one of the subspecies of Calandrella blanfordi (formerly treated as subspecies of C. cinerea).

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Old Friday 23rd June 2017, 09:12   #15
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... seems to be a synonym for one of the subspecies of Calandrella blanfordi (formerly treated as subspecies of C. cinerea).
IOC (7.01):
Quote:
Calandrella blanfordi (Shelley, 1902)
C. b. eremica (Reichenow & Peters, JL, 1932)
C. b. blanfordi (Shelley, 1902)
C. b. daaroodensis White, CMN, 1960
If so; why isn´t Rüppel's name (of 1835) valid?
---

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Old Friday 23rd June 2017, 09:19   #16
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Sclater 1930 (Syst. Av. Ethiop.: 333 [here] -- not sure full view access will be granted to everyone in the world) has a footnote to his Tephrocorys cinerea ruficeps (Rüpp.) that reads:
Quote:
Tephrocorys cinerea erlangeri Neumann, Journ. Orn. 1906, p. 239 : Sheikh Mohamed, on the Webbi River, does not appear to be separable.
Alauda ruficeps Rüppell 1835 [OD] is preoccupied by Alauda arvensis ruficeps Bechstein 1795 [OD], and thus cannot be used.
Presumably erlangeri Neumann was picked up to replace it when this was realized?

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Old Friday 23rd June 2017, 09:30   #17
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ruficeps Rüppell is in the synonymy of erlangeri Neumann in the Peters Check-list: [here].

Last edited by l_raty : Friday 23rd June 2017 at 10:26. Reason: not 'Naumann'
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Old Friday 23rd June 2017, 09:32   #18
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The main issue (at least for me) is to understand which one of today's taxon Dr. Blanford regarded as different from the Lark that since 1902 have his name.

Was I thereby correct in assuming that he regarded it as distinct from today's Erlanger's Lark ... ?

The taxonomy of those smaller Larks are sure tricky (more than tricky) to follow!
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Old Friday 23rd June 2017, 09:36   #19
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Thanks, Laurent!

You answer and reply even quicker than I can write!
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Old Friday 23rd June 2017, 15:29   #20
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Calandrella.
I am returning to this thread very briefly to highlight the differences that become apparent from using the varied existing checklists. I prefer paper, and cannot find my way through the maze that is the IOC World Bird List, with its Life Lists, Master Lists, Multilinguals, Comparisons, etc, etc!. My own world checklist is the splitter's paradise of a heavily annotated copy of Howard & Moore ed. 3, which I can edit in the twinkling of an eye (I don't yet have the courage to write in ed. 4!), but for this Forum, and, obviously, my Key, I use the Illustrated Checklist, with its taxonomy, maps and illustrations, as my base. In vol. 2 of the latter, Blanford's Lark Calandrella blanfordi contains only two subspecies, blanfordi and erlangeri (whence #14). There also the Rufous-capped Lark C. eremica contains two subspecies, eremica and daaroodensis. Howard & Moore ed. 4, treat all four taxa as subspecies of C. blanfordi. I assume the IOC List treats erlangeri as a separate species. Perhaps in the future, to avoid confusion, we should state which list/system is being followed.
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Old Friday 23rd June 2017, 16:36   #21
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Calandrella.
I assume the IOC List treats erlangeri as a separate species. Perhaps in the future, to avoid confusion, we should state which list/system is being followed.
Yes; IOC splits erlangeri, but lumps eremica into blanfordi.

All those various IOC lists are just different formats for the same list. Mostly you just want the "Master list" to peruse their taxonomy, or the "Life list+" if you want to record into a spreadsheet (though IOC changes fast enough - four times a year - that I'd recommend software that can handle the updates rather than manual spreadsheet entry).
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Old Friday 23rd June 2017, 17:04   #22
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Yes; IOC splits erlangeri, but lumps eremica into blanfordi.
...which used to be a rather widespread treatment -- Erlanger's Lark, monotypic; Blanford's Lark, with sspp blanfordi, eremica and daroodensis. (It's also the treatment that was adopted in the paper edition of HBW.)

blanfordi being close to erlangeri rather than to the other two, emerged as a surprise from the results of Stervander et al 2016 ([pdf here]; see posts #3 et seq. above). Interestingly, the text of the Illustrated Checklist doesn't even cite Stervander et al., and suggests that the BirdLife team reached the same conclusion based on morphology.
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Old Friday 23rd June 2017, 21:27   #23
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ruficeps Rüppell is in the synonymy of erlangeri Neumann in the Peters Check-list: [here].
Friedmann in 1932 recognises the problem with A. ruficeps and names these birds for Fuertes.
http://www.biodiversitylibrary.org/p...e/185/mode/1up .
But then in 1937 reject that new name and calls them A. erlangeri. Page 38 of BIRDS COLLECTED BY THE CHILDS FRICK EXPEDITION TO ETHIOPIA:
http://www.biodiversitylibrary.org/p...ge/62/mode/1up .
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Old Saturday 24th June 2017, 07:25   #24
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Thanks guys, for all your help sorting out this query!

Well, I´m content: Thereby that certain "T. ruficeps", mentioned by Shelley in 1902, was/is the equal of today's erlangeri NEUMANN 1906.

Dr. Blanford sure must have been quite a sharp-eyed birder in those days.

"T. ruficeps" ... over and out!
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Old Monday 2nd July 2018, 19:12   #25
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Calandrella eremica, Calandrella erlangeri

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Martin Stervander, Per Alström, Urban Olsson, Ulf Ottosson, Bengt Hansson, Staffan Bensch. Multiple instances of paraphyletic species and cryptic taxa revealed by mitochondrial and nuclear RAD data for Calandrella larks (Aves: Alaudidae). Molecular Phylogenetics and Evolution. In Press, Accepted Manuscript, Available online 26 May 2016.

[abstract]
IOC Updates Diary July 2

Accept split of Rufous-capped Lark

Treat Erlanger’s Lark as subspecies of Blanford’s Lark
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