Implications(long!) of AOU Supplement for other lists

Subject: Implications(long!) of AOU Supplement for other lists
From: John Penhallurick <>
Date: Sun, 27 Jul 1997 16:33:34 +1000
What follows deals particularly with implications for the Australian list,
but the remarks on Swiftlets and Tits will be relevant to many lists.

Implications of the AOU Checklist Supplement , "Forty-First Supplement to
the American Ornithologists' Union Checklist of North American Birds" Auk,
114(3):542-552, for the Australian List.

The changes to the AOU List mainly concern North American and South American
Species.  Howver the taxonomic basis of some decisions has relevance to some
species on the Australian list.

 p.543. The albatrosses, family Diomedeidae, are arranged in four genera,
following the results of genetic studies by Nunn[
Nunn,Cooper,Jouventin,Robertson & Robertson,1996, 'Evolutionary
relationships among extant albatrosses (Procellariiformes: Diomedeidae)
established from complete cytochrome-b gene sequences' Auk:113:784-801.]
The sequence and new name combinations are Diomedea exulans Linnaeus,
Phoebastria nigripes (Audubon), Phoebastria immutabilis  (Rothschild),
Phoebastria albatrus (Pallas),Thalassarche chlororhynchus  (Gmelin),
Thalassarche cauta (Gould), Thalassarche melanophris (Temminck), Phoebetria
palpebrata (Forster). English names do not change.

Basically, the four groups each group closely together in genetic terms, and
the genetic differences between each of the four groups are greater than
those between other groups that have traditionally been reconised as
different species.

If the arbiters of the Australian Checklist accept this, and I think they
should, the world list will become:

*Diomedea exulans
Diomedea amsterdamensis
*Diomedea epomophora

Phoebastria irrorata (cf AOU checklist p.549)
Phoebastria albatrus
Phoebastria nigripes
Phoebastria immutabilis

*Thalassarche melanophris
*Thalassarche cauta 
*Thalassarche chrysostoma  (cf AOU checklist p.549)
*Thalassarche chlororhynchus
*Thalassarche bulleri

*Phoebetria fusca
*Phoebetria palpebrata

I have placed an asterisk (*) against species on the Australian list.

'p.166.The specific name of the American Golden-Plover, erroneously changed
to dominicus in the 40th Supplement, is corrected to the adjectival form

'p.322. The genus Aerodramus Oberholser replaces Collocalia G.R.Gray,
inserted in the main list by the 40th Supplement, following Lee et al.
(1995). Aerodramus bartschi, Guam Swiftlet, replaces Collocalia bartschi ...

Lee,Clayton, Griffiths & Page, 1996, 'Does behavior reflect phylogeny in
swiftlets (Aves:Apodidae)? A test using cytochrome b mitochonrial DNA
sequences' Proc.Nat.Acad.Sci USA,93: 701-7096.  
found that all the echolocating species they analysed ( namely spodiopygius,
sawtelli,bartschi,salanganus, fuciphagus, elaphrus,
francicus,brevirostris,maximus, terraereginae AND Hydrochous gigas group

Our discussion relates to Bootstrap consensus tree in Fig.3, p. 7094.

The authors state of this: "The basal nodes corresponding to the
Apodidae(95%), the Cypseloidinae(82%) [including Steptoprocne] and the
Apodinae(96%) all have good support, as does the genus Collocalia(85%) [ie.
Collocalia in the narrow sense, as opposed to Aerodramus]. However,some
relationships in the Apodini,Chaeturini and remaining Collocaliini are less
robust.  Examination of the bootstrap trees revealed that much of this
ambiguity was due to uncertainty in the placement of Apus apus, Apus
nipalensis, Cypsiurus and Chaetura.  These taxa are unstable and tend to
'wander' over the tree, reducing bootstrap values for nodes that might
otherwise have good support."

Collocalia troglodytes,linchi and esculenta ie. the non-echo-locating
swiftlets form a well-supported group, that is somewhat remote from the
Aerodramus group.

The Aerodramus group is also well supported.

These data suggest something like the following, allowing for the
uncertainty attaching to Cypsiurus and Chaetura:

Family                          APODIDAE

Subfamilies     Cypseloidinae                   Apodinae

Tribes                                  Collocaliini           Apodini          

Subtribes                                       Cypsiurina Chaeturina 
Aerodramina Hydrochous

Some uncertainty attaches to Hydrochous.  It is clearly related to
Aerodramus, but one cannot with certainty say it is a sister-group for
Aerodramus.  It seems best at present to retain the genus Hydrochous and
place it next to Aerodramus in the sequence of genera.

Some implications at the level of species are interesting. 

Aerodramus bartschi forms a well-supported sister-group(75%) with
A.sawtelli, usually seen as a member of the leucophaeus superspecies..
Unfortunately, A. vanikorensis, with which bartschi has sometime been
merged, was not studied, nor were either A.leucophaeus or A.ocistus. But
A.salanganus, supposedly a member of the vanikorenis superspecies is not a
sister species of bartschi,but a member of a group which is a sister-group
to the group containing bartschi.  It looks as if bartschi's species status
is confirmed, contra Sibley and Monroe,Supplement,1993,p.26..   

A.terraereginae forms a fairly well-supported sister-species(61%) to a group
formed by the combination of A.brevirostris and A.maximus.  The connection
of A.terraereginae with any forms of spodiopygius is remote.
A.terraereginae should be recognised as a good species. 

A.elaphrus and A.francicus form as sister-group, confirming the superspecies
relationship in Sibley and Monroe (1990:134)

A further really interesting result is that several "species" are reliably
revealed to be polyphyletic.  
A.spodiopygius assimilis from Vanua Balavu is not close to A.spodiopygius
assimilis from Suva, and neither is a sister group to any other form of
A.spodiopygius.  Similarly  A.salanganus from Balambangan was not a sister
group to A.salanganus from Gomantong Caves.  The same is true of two
specimens of A.fuciphaga vestitus from the same location!

The authors comment: 'Given the instability of swiftlet species and
subspecies boundaries, this[polyphiletic character of several species] is
not entirely surprising, although it is possible that the discordance
reflects lineage sorting or past hybridization events.' (p.7094).

I would say that Fig 3 strongly supports the awarding of species status to
at least A.sawtelli, A.bartschi, A.elaphrus, A.francicus, A.germani,
A.brevirostris, A maximus and A.terraereginae.

Another feature of the new AOU list, which, although it does not effect the
Australian list, affects many other lists, is its acceptance of the
conclusions of Slikas, Sheldon & Gill, 1996,Phylogeny of titmice (Paridae):
Estimate of relationships among subgenera based on DNA-DNA hybridisation'
Journ.Avian Biol.,27:70-82.  This study, which used complete matrices, found
essentially that the genetic differences between subgenera put forward by
Hellmayr, 1903 (Paridae,Sittidae u.Certhiidae.Das Tierreich,no.18) and
discussed by Thielcke,1968,Beiheft der Vogelwelt,1:147-164, and more
accessibly by Hailman,1989,Wilson Bull.,101:305-343, were greater than those
between some other genera.  As the authors note: 'The distances between
parid subgenera are similar in magnitude to those distinguishing genera of
herons (excluding two clades, Botaurus and Ixobrychus,Cochlearius and
Tigrisoma) and of swallows, but less than those distinguishing most genera
of ducks, hummingbirds, and emberizids."

The AOU has taken this to its logical conclusion and recognised six genera
in place of Parus.
This leads to the following reassignments, which include not only taxa
analysed, but forms recognisably very close to them.

Poecile palustris       Marsh Tit
Poecile lugubris        Sombre Tit
Poecile montanus        Willow Tit
Poecile carolinensis    Carolina Chickadee
Poecile atricapillus    Black-capped Chickadee
Poecile gambeli         Mountain Chickadee
Poecile sclateri        Mexican Chickadee
Poecile superciliosus   White-browed Tit
Poecile davidi          Rusty-breasted Tit
Poecile cinctus         Siberian Tit
Poecile hudsonicus      Boreal Chickadee
Poecile rufescens       Chestnut-backed Chickadee

Periparus rufonachalis  Dark-grey Tit
Periparus rubidiventris Rufous-vented Tit
Periparus melanolophus  Black-crested Tit
Periparus ater          Coal Tit

Lophophanes cristatus   Crested Tit
Lophophanes dichrous    Grey-crested Tit

Parus major             Great Tit
Parus bokharensis       Turkestan Tit

Cyanistes cyanus        Azure Tit
Cyanistes flavipectus   Yellow-breasted Tit

Baeolophus wollweberi   Bridled Titmouse
Baeolophus inornatus    Oak Titmouse
Baeolophus ridgwayi     Juniper Titmouse
Baeolophus bicolor      Tufted Titmouse 
Baeolophus atricristatus        Black-crested Titmouse. (This is not yet 
as a species distinct from bicolor by the AOU, but its recongition is, in my
view, inevitable.)

There are problems, however, in applying the family reanalysis to members of
the genus Parus that did not form part of the taxa analysed.  In some cases,
one can classify taxa on the basis of their close similarity to taxa that
were analysed, as has been done above.  But until analysis is carried out on
some intermediate forms, one can not be sure which genus they belong to.  So
in my list, I have reassigned members of Parus to the new genera where this
could be done with confidence, and left others with Parus as their genus.
However, this raised another problem.  A basic principle of taxonomy is that
members of the same genus should appear together in the list.  But Parus
remains one of the six new genera.  And if we grouped together all species
with Parus as genus, we would be mixing up members of the new, more
restricted genus (which is related to Poecile, Baeolophus, Cyanistes etc.)
with members of the old, inclusive genus Parus.  Moreover the order in the
list represents the best guess we have at present as to what the
relationships of unanalysed forms in the list are to reassigned forms.  So,
in the knowledge, or hope, that these uncertain forms will shortly be
reassigned to the new genera, I have left them in their place, which means
that you will see specimens with the genus Parus mixed up with the names of
other genera.  The only species that can with certainty be assigned to the
new, restricted definition of Parus are Parus major, Great Tit, and Parus
bokharensis, Turkestan Tit.  So all other instances of Parus represent forms
to be reassigned to the new genera after they are analysed.

John Penhallurick

Associate Professor John M. Penhallurick<>
Canberra, Australia
Phone BH( 61 6) 201 2346   AH (61 6 2585428)
FAX (61 6) 258 0426
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                University of Canberra, PO Box 1, BELCONNEN, A.C.T.2616,
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                "I'd rather be birding!" 

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