Proposals (220) to South American Classification Committee
Merge Nesomimus
into Mimus
Background: The mockingbirds of the Galapagos Islands
have traditionally been placed in their own genus, Nesomimus, based on
plumage and morphology, although the rationale for recognition of Nesomimus
has been questioned by some authors (see references in Arbogast et al. 2006).
New information: Arbogast et al. (2006) sequenced
> 2650 bp of 4 mtDNA genes from Nesomimus populations from 11 islands
as well as all members of the genus Mimus. Their molecular phylogeny
demonstrates that Nesomimus is monophyletic. Their phylogeny suggests
that it is embedded in Mimus, and that its closest relatives are,
somewhat surprisingly in terms of plumage, probably the northern group (M.
gundlachii, M. polyglottos, M. gilvus). [Of biogeographic
interest is that several Galapagos taxa also have their closest relatives in
the West Indies, and the northern Mimus clearly have the better track
record as dispersers and colonists than the South American group.] [Also of
interest is a problem in species limits within the Galapagos, but that's a
separate proposal.]
Analysis: Although Arbogast et al. (2006) word their findings in a way
that suggests that there is no doubt that Nesomimus is embedded in Mimus,
a closer look shows that the support for the critical nodes is below the
thresholds usually required for treating the findings as beyond a reasonable
doubt. Specifically, the support level for the node that links Nesomimus
with northern Mimus is only 49% (bootstrap probability). As my colleague
Robb Brumfield noted, it would be interesting to do a Shimodaira-Hasegawa test
to see if there's a significant difference between the trees they present and a
tree in which Mimus is constrained to be monophyletic.
Nonetheless, I'm just quibbling. I suspect that additional
sampling will elevate the support for the critical nodes. It seems not only
sensible but also inevitable that Nesomimus is embedded in Mimus,
i.e., derived from extant species or ancestral species within Mimus rather
than from a shared common ancestor outside Mimus. In other words, it
seems exceptionally unlikely that Nesomimus was derived from the shared
ancestor of all extant Mimus, and even if it were, ranking it as a
separate genus would boil down to a matter of taste. Even if Nesomimus is
not embedded within extant Mimus, it is more closely related to Mimus
than to the sister taxon to all mockingbirds, Oreoscoptes; that node IS
strongly supported. Therefore, I would support the merger of Nesomimus into
Mimus.
Recommendation: YES. Our current sequence does not reflect
phylogenetic hypotheses. Although linear sequences are a poor way to portray
relationships, for consistency we should make those sequences as consistent as
possible with available data, and those data suggest that Nesomimus is
embedded within or, at worst, the sister group to, all Mimus. To address
the uncertainty for the support for the Nesomimus-northern
Mimus sister relationship, I suggest we move the Galapagos species to an
intermediate position between northern and southern Mimus, with a Note
to explain our caution. Thus, our sequence would place the Galapagos species
between M. gilvus and M. thenca in the current sequence.
Literature Cited (see SACC Biblio for the rest):
ARBOGAST, B., S. V. DROVETSKI, R. L. CURRY, P. T. BOAG, G. SEUTIN,
P. R. GRANT, B. R. GRANT, AND D. J. ANDERSON. 2006. The origin and
diversification of Galapagos mockingbirds. Evolution 60: 370-382.
V. Remsen,
June 2006
Note added 8/22/07: See also Lovette & Rubenstein
(2007), although as Daniel pointed out to me, they used mostly previously
sequenced material: (p. 1035: "Some or all DNA sequences from 16 Mimidae
taxa ... were derived from previous studies [Hunt et al. 2001, Barber et al.
2004, Arbogast et al. 2006])"
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Comments from Jaramillo: "YES - Data are solid, and
in life Nesomimus is not that different than let's say a Bahama
Mockingbird, so this is not surprising. Van's point about how sensible it is
that the Galapagos taxa would be expected to be imbedded in the genus, rather
than an offshoot from a nearest relative of the mainland genus is a good one.
This logic can be applied to many island taxa; the interesting situation is
when the relationships do not fit this expectation (Caribbean trogons, todies
etc.)."
Comments from Stiles: "YES. This arrangement might
not be as surprising as it seems given the probable tectonic history of the
Caribbean plate, which as I understand it moved from west to east through the
gap between North and South America well before the final closure of this gap
with the isthmus of Panama, producing and pushing island arcs ahead of it as it
moved. Thus, I don't find it so terribly surprising that a Caribbean genus like
Spindalis turns out to be sister to the rest of the extant tanagers (Burns?):
it probably represents an old lineage present at the start of the
differentiation of the tanagers that hopped aboard such an island arc as it
passed through, remained isolated and suffered relatively limited
differentiation in the Antilles thereafter, while the rest of the lineage
either went extinct or differentiated much more explosively on the mainland(s)
what with orogenies, sea level changes, glaciations and climatic changes, etc.
not to mention the wealth of interspecific interactions (or alternatively, it
evolved on an early island arc and hopped off onto the mainland as it passed
by; subsequent differentiation on the mainland was explosive, on the islands
very minor. The presence of related forms in the Galapagos and the Antilles
could mean that representatives of some other groups (e. g., Darwin's finches
got off the boat in the other direction at a relatively early date. Hence, the
conclusion that "Nesomimus" are closer to
northern-Antillean mockingbirds than to southern ones might not be wholly
unique or unprecedented (unless I'm speculating overly wildly??)."
Comments from Robbins: "NO. The molecular data
unequivocally demonstrate that Nesomimus is monophyletic; however,
whether it is embedded within Mimus is open to question (note the poor
nodal support) and even the sister taxon to Nesomimus is
equivocal. Although including Nesomimus within Mimus likely will
prove correct, the current data set does not establish that as fact. Before
speculating about historical biogeographical scenarios, we should first solidly
establish relationships and be consistent in our application of minimum
criteria for evaluating those data."
Comments solicited from
Peter Grant: "I do not have a
strong opinion about the question of the appropriate genus. I tend to be
conservative, and do not want to change a name unless the case is very clear.
As Robbins indicates the critical node is not well supported, and if it were
left to me I would not change the genus name until that node is resolved. I
must admit that I expect Nesomimus to fall within Mimus.
Another reason for being conservative is that I don’t like pinning a judgment
on just one (mitochondrial) molecule, even when several genes have been assayed
for greater statistical resolution. There are plenty of instances elsewhere of
strong conflict between the signals of relatedness from nuclear and
mitochondrial genes."
Comments from Pacheco: "NO. Em vista das opiniões sobretudo de Mark Robbins e
Peter Grant, voto pela manutenção de Nesomimus, até que mais estudos
complementem as sugestões ora levantadas e resolvam a questão."