Abstract.---The unwitting inclusion of convergent characters in phylogenetic estimates
poses a serious problem for efforts to recover phylogeny. Convergence is not inscrutable,
however, particularly when one group of characters tracks phylogeny and another set tracks
adaptive history. In such cases, convergent characters may be correlated with one or a few
functional anatomical units and readily identifiable using comparative methods. Stifftail
ducks (Oxyurinae) offer one such opportunity to study correlated character evolution and
function in the context of phylogenetic reconstruction. Morphological analyses place
stifftail ducks as part of a large clade of diving ducks that includes the sea ducks
(Mergini), Hymenolaimus, Merganetta, and Tachyeres, and possibly the pochards (Aythyini).
Molecular analyses, on the other hand, place stifftails far from other diving ducks and
suggest, moreover, that stifftails are polyphyletic. Mitochondrial cytochrome b gene
sequences of eight stifftail species traditionally supposed to form a clade were compared
with each other and with sequences from 50 other anseriform and galliform species.
Stifftail ducks are not the sister group of sea ducks, but lie outside the typical ducks
(Anatinae). Of the four traditional stifftail genera, monophyly of Oxyura and its sister
group relationship with Nomonyx are strongly supported. Heteronetta probably is the sister
group of that clade, but support is weak. Biziura is not a true stifftail. Within Oxyura,
Old World species (O. australis, O. leucocephala, O. maccoa) appear to form a clade, with
New World species (O. jamaicensis, O. vittata) branching basally. Incongruence between
molecules and morphology is interpreted to be the result of adaptive specialization and
functional convergence in the hind limbs of Biziura and true stifftails. When morphological
characters are divided into classes, only hind-limb characters are significantly in conflict
with the molecular tree. Null models of synonomous and nonsynonomous substitution based on
patterns of codon-degeneracy and chemical dissimilarity, likewise, indicate that the
nucleotide and amino acid changes postulated by the molecular tree are more plausible than
those postulated by the morphological tree. These findings teach general lessons about the
utility of highly adaptive characters (in particular those related to foraging ecology) and
underscore the problems that convergence can pose for attempts to recover phylogeny. They
also demonstrate how the concept of natural data partitions and simple models of evolution
(e.g., parsimony, likelihood, neutrality) can be used to test the accuracy of independent
phylogenetic estimates and provide arguments in favor of one tree topology over another.
[Anseriformes; Anatidae; Oxyurinae; cytochrome b; congruence analysis; functional morphology;
behavior, diving, foraging ecology.]
Abstract.---There are 14 species of marmots distributed across the Holarctic, and despite
extensive systematic study, their phylogenetic relationships remain largely unresolved. In
particular, comprehensive studies have been lacking. A well-supported phylogeny is needed to
place the numerous ecological and behavioral studies on marmots in an evolutionary context.
To address this situation, we obtained complete cytochrome (cyt) b sequences for 13 of the
species and partial sequence for the 14th. We employed a statistical approach to both
phylogeny estimation and hypothesis testing using parsimony and maximum likelihood based
methods. We conducted statistical tests on a suite of previously proposed hypotheses of
phylogenetic relationships and biogeographic histories. The cyt b data strongly support the
monophyly of Marmota and a western montane clade in the Nearctic. The results are consistent
with an initial diversification in North America followed by an invasion and subsequent
rapid diversification in the Palearctic. These analyses reject the two major competing
hypotheses of M. broweri’s phylogenetic relationships: namely, that it is the sister
species to camtschatica of eastern Siberia and that it is related closely to caligata of the
Nearctic. The Alaskan distribution of M. broweri is best explained as a reinvasion from the
Palearctic but a Nearctic origin can not be rejected. Several other conventionally recognized
species groups can also be rejected. Social evolution has been homoplastic, with large
colonial systems evolving in two groups convergently. The cyt b data do not provide
unambiguous resolution of several basal nodes in the Palearctic radiation, leaving some
aspects of pelage and karyotypic evolution equivocal.
[Marmota; cyt b; phylogenetics; Holarctic; Beringia; hypothesis testing]
Abstract.---Cladistic analyses of the nuclear ribosomal DNA (nrDNA) internal transcribed
spacer (ITS) sequences from 55 samples corresponding to 34 taxa in the genus Armeria reveal
that ITS sequence diversity among and within species utterly conflicts with patterns of
morphological similarity. Three facts are apparent from the results here reported: (1)
different samples of a single subspecies, A. villosa subsp. longiaristata, appear in three
of the five major clades; (2) samples of at least one of the six subspecies of A. villosa
appear in four of the five major clades; and (3) composition of major clades shows greater
congruence with the geographic origin of plants than with the traditional systematic
arrangement based primarily on morphology. Specifically, the clades here termed Ia, II, III,
and IV each encompass terminals restricted to geographically delimited areas. There are
alternative explanations for the ITS pattern, but the most likely one is that nucleotide
positions supporting the major clades are due, in some of the samples, to concerted evolution
following horizontal transfer (gene flow) rather than to recency of common ancestry. This
interpretation is consistent with previous systematic and experimental evidence and implies
that reticulation in Armeria may be extensive. Harlan and de Wet (1963, Evolution 17:497-501)
proposed the compilospecies concept to account for situations in which a genetically
"aggressive" species captures portions of the genome of other sympatric species via extensive
introgression. Evidence of extensive reticulation, ecological diversification, and
geographic pattern indicates that A. villosa may fit the compilospecies concept, which is
here supported on molecular grounds for the first time.
[Armeria; compilospecies; concerted evolution; hybridization; Plumbaginaceae; ribosomal DNA;
reticulate evolution]
Abstract.---Cladistic relationships between 7 parts of the hesionid polychaete group
Heteropodarke are assessed in a parsimony analysis based on 34 morphological characters.
Taxon names are defined by apomorphy-based phylogenetic definitions, without reference to
Linnean ranks or types. Species entities are omitted and denied any role in taxonomy; taxon
names refer to monophyletic groups only. Linnean binomial species names are not employed,
and all taxa are assigned uninomials. Previously known parts of Heteropodarke (Africana
Hartmann-Schršder, 1974; Formalis Perkins, 1984; Heteromorpha Hartmann-Schršder 1962;
Lyonsi Perkins, 1984; Xiamenensis Ding, Wu, and Eestheide, 1997) are reexamined, and
Bidentata, new taxon, and "Zmyrina" (informal name) are described from Papua New Guinea and
Belize, respectively. The new taxon Crassichaetae is named for a subgroup of
Heteropodarke, which is diagnosed by enlarged, anteriorly situated falcigers, and includes
Africana, Heteromorpha, Lyonsi, and Xiamenensis. Within this group Africana and Heteromorpha
are treated as taxa inquirendae.
[Apomorphy-based definitions; Linnean nomenclature; phylogenetic taxonomy; polychaetes;
ranks; species concepts; species names; types.]
Abstract.---Linnaean binomial nomenclature is logically incompatible with the phylogenetic nomenclature of de Queiroz and Gauthier (1992, Annu. Rev. Ecol. Syst. 23:449-480), because the former is based on the concept of genus, thus making this rank mandatory, while the latter is based on phylogenetic definitions and requires the abandonment of mandatory ranks. Thus, if species are to receive names under phylogenetic nomenclature, a different method must be devised to name them. Thirteen methods for naming species in the context of phylogenetic nomenclature are contrasted with each other and with Linnaean binomials. A fundamental dichotomy among the proposed methods distinguishes those that retain the entire binomial of a preexisting species name from those that retain only the specific epithet. Other relevant issues include the stability, uniqueness, and ease of pronunciation of species names, their capacity to convey phylogenetic information, and the distinguishability of species names governed by a code of phylogenetic nomenclature both from clade names and from species names governed by the current codes. No method is ideal. Each has advantages and drawbacks, and preference for one option over another will be influenced by one's evaluation of the relative importance of the pros and cons. Moreover, sometimes the same feature is viewed as an advantage by some and a drawback by others. Nevertheless, all of the proposed methods for naming species in the context of phylogenetic nomenclature provide names that are more stable than Linnaean binomials.