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Journal ArticleDOI

Evolutionary Relationships among the North American Mallards

01 Jan 1961-The Auk (Oxford University Press (OUP))-Vol. 78, Iss: 1, pp 3-43
TL;DR: An attempt to understand the evolutionary relationships existing within a group of mallardlike ducks native to North America.
Abstract: THIS study is the report of an attempt to understand the evolutionary relationships existing within a group of mallardlike ducks native to North America. The group includes the Common Mallard, Anas platyrhynchos platyrhynchos L.; the Black Duck, Anas rubripes Brewster; the Florida Duck, Anas fulvigula fulvigula Ridgway; the Mottled Duck, Anas fulvigitla niiaculosa Sennett; the Mexican Duck, Anas diazi diazi Ridgway; and the New Mexican duck, Anas diazi novimexicana Huber. All but one of these (the Common Mallard) are restricted to North America, and all these American forms possess a sexually nondimorphic plumage. In all other respects they are extremely similar to the Common Mallard, and a study of their relationships to this form was believed possibly to provide an instructive example of speciation.

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Summary

  • Library preparation was performed using the low-input genomic 80 DNA sequencing kit protocol for SQK-MAP006 per manufacturer’s instructions.
  • Taxonomic analysis was performed with Kraken and plotted using Krona [6].
  • The authors detected VRE (E. 92 faecium and/or E. faecalis), vanA and/or vanB in all samples, and predominant CRO organisms 93 (E. cloacae, P. aeruginosa and K. pneumoniae) in 3 samples (Figure 1A-B).
  • Both platforms 94 detected the same or similar species as the top three dominant organisms (Table 1). 95 was not certified by peer review) is the author/funder.
  • Swabs #2 and 101 #10 had vanB detected and Swab #7 had blaKPC detected, corresponding to culture and PCR 102 results from the rectal swab.
  • Newer 142 sequencing methods (rapid extraction and library kits, Flongle) and flow cells (MinION R.9 and 143 R.10) from Oxford Nanopore may improve accuracy significantly [15].
  • (I) Resistome analyses demonstrated the presence of both vancomycin (vanA 252 operon) and carbapenem resistance genes based on adaptation of using ResFinder and additional BLAST analyses 253 254 w as not certified by peer review ) is the author/funder.

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University of Nebraska - Lincoln University of Nebraska - Lincoln
DigitalCommons@University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln
Papers in Ornithology Papers in the Biological Sciences
1961
Evolutionary Relationships among the North American Mallards Evolutionary Relationships among the North American Mallards
Paul A. Johnsgard
University of Nebraska-Lincoln
, pajohnsgard@gmail.com
Follow this and additional works at: https://digitalcommons.unl.edu/biosciornithology
Part of the Ornithology Commons
Johnsgard, Paul A., "Evolutionary Relationships among the North American Mallards" (1961).
Papers in
Ornithology
. 62.
https://digitalcommons.unl.edu/biosciornithology/62
This Article is brought to you for free and open access by the Papers in the Biological Sciences at
DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in Ornithology by an
authorized administrator of DigitalCommons@University of Nebraska - Lincoln.

EVOLUTIONARY RELATIONSHIPS AMONG THE
NORTH AMERICAN MALLARDS
PAUL A. Jo•X•SG.•
This study is the report of an attempt to understand the evolutionary
relationships existing within a group of mallardlike ducks native to North
America. The group includes the Common Mallard, `4nas platyrhynchos
platyrhynchos L.; the Black Duck, `4nas rubripes Brewster; the Florida
Duck, `4nas fulvigula fulvigula Ridgway; the Mottled Duck, .4nas fulvi-
gula macuIosa Sennett; the Mexican Duck, .4nas diazi diaxi Ridgway;
and the New Mexican duck, .4nas diazi novimexicana Huber. All but
one of these (the Common Mallard) are restricted to North America,
and all these American forms possess a sexually nondimorphic plumage.
In all other respects they are extremely similar to the Common Mallard,
and a study of their relationships to this form was believed possibly to
provide an instructive example of speciation.
REVIEW OF RANGES, POPULATIONS, AND NOMENCLATURE
The Mexican and New Mexican Ducks
In 1886 Ridgway described a new species of mallard (.4nas diazi)
from Puebla, Mexico, which, similar to the then recently (1874) de-
scribed Florida Duck, differed from the Common Mallard (`4nas platy-
rhynchos) in its lack of sexual dimorphism. According to Ridgway, the
new species differed from the Florida Duck in its more Mallardlike
characteristics, namely the distinct band of white on the secondary wing
coverts and its less-fulvous over-all coloration. Thirty-six years later,
Huber (1920) described another species of mallard from New Mexico,
which he named `4nas novimexicana. Huber was seemingly unaware of
the similarities shown by the New Mexican Duck to the Mexican Duck,
but in 1922 Conover referred to an extralimital Nebraskan specimen as
.4has dia•i novimexicana, and Phillips (1924) treated the New Mexican
Duck in the same manner in his monograph. Although the close rela-
tionship between the New Mexican and the Mexican ducks was there-
fore realized relatively early, the paucity of specimens and field observa-
tions made the exact geographic relationship between the two forms
uncertain. In 1946 Lindsey summarized the situation as follows: "The
known nesting range of the New Mexican Duck is confined to a small
area of the south-western United States, but the presumptive range
extends southwards into Chihuahua, Mexico, where its relation to the
northern breeding limits of the Mexican Duck (.4. diazi diazi) is unde-
terrained."
Published in the Auk (1961) 78. Copyright 1961, University of California. Used by permission.
http://elibrary.unm.edu/sora

4 J Olt NSGARD, Relationships among Mallards [ Auk
ß I. Vol. 78
ARIZONA i 4G : NEW MEXICO
! ' :8 4G TEXAS
........
j .
.... , •, '•,
',
SINALOA )---" ,[ ( '
'•. /•--- •.-. ,.•-
"-•'-, ..• ,/ '• ,,
) ß ( ,' :, '.,
,• ZACATECAS ,-"' ,
Sight Specimen Range
M ex•c•In Duck O ß ,.r-•
(inc) New Mexicofi")
Mottled Duck O ß __
O 50 I00 I•0 20<) •0
JALISCO •.
OL So•ulo ?
MICNOACAN
An attempt has been made to map the geographic ranges of the
Mexican and New Mexican ducks, and thus to help visualize their geo-
graphic relationships. This was done by plotting all available specimen
records and such sight records as appeared justified from the literature,
personal communications, and the major United States collections. The
Published in the Auk (1961) 78. Copyright 1961, University of California. Used by permission.
http://elibrary.unm.edu/sora

Auk 1 Jon•-sg^Ru, Relationships among Mallards 5
Vol. 78 J
resulting map (Figure 1) shows fairly close agreement with thc com-
parable maps of Dclacour (1956), Arcllano and Rojas (1956), and
Leopold (1959). It follows the map of Leopold in indicating an unbroken
range bct;vccn the New Mexican and Mexican populations, although
records arc spotty for the area in question. If birds at the northern parts
of the range arc migratory, for which there is some evidence, it is probable
that they winter in the heart of the range of diazi; habitat conditions arc
very poor on the Chihuahua lakes, and they arc unattractive to waterfowl
(Saunders and Saunders, 1949).
Of all the races of North American mallards, perhaps least is known
about the Mexican Duck, and this is certainly the case regarding the
total population size. Since 1947 the U.S. Fish and Wildlife Service
has surveyed the lakes of interior Mexico during its annual January
waterfowl inventories. Mr. R. H. Smith, who has flown this route each
year since 1951, has provided (in litt.) the data obtained for the Mexican
Duck during these years. The sinailest total number of Mexican Ducks
recorded was 780 in 1951, and the largest total was 10,322 in 1958.
Although the northern Chihuahuan lakes--Toronto, Bustillos, and Mexi-
canos--•vere surveyed, it is of interest that no Mexican Ducks have been
recorded there; Lago Santiaguillo, Durango, is apparently the northern
wintering limit of the Mexican Duck in Mexico. Its western limit would
appear to be Lago Sayula, Jalisco, and the eastern limit in the 1natshy
region near Oriental, Puebla. These data would indicate that the size
of the adult Mexican Duck population is probably less than 20,000 birds.
New Mexican game biologists have estimated that the population in their
state is extremely snmll, possibly not over 100 adult birds.
The Florida and Mottled Ducks
In 1874 Ridgway described a new form of mallard from St. Johns,
Florida, which he considered to be a subspecies of the Black Duck. As
the Black Duck was at that time called Anas obscura, the new bird was
named Anas obscura var. fulvigula. It differed mainly from the Black
Duck in that the malar region, chin, and throat were immaculate buff
rather than streaked with dusky, and the plumage had a general
ochraceous, rather than dusky, tone. At the time it was known only
from Florida. Later (1878) Ridgway relegated it to the rank of a full
species, Arias fulvigula. Eleven years later, Sennerr (1889) described
from Texas yet another species of mallard, which he called .4has
maculosa. Sennett stated that the most marked differences between
zt. )•ulvigula and A. maculosa were the latter's streaked, rather than plain
buff, cheeks and a green, rather than purple, speculum in addition to a
mottled, rather than streaked, general coloration. The Mottled Duck
Published in the Auk (1961) 78. Copyright 1961, University of California. Used by permission.
http://elibrary.unm.edu/sora

6 Joa•s•^RD, Relationships among Mallards [ .•.uk
[Vol. 78
was added to the A.O.U. Check-list in 1890 as Anas Iulvigula •naculosa,
and no later arguments for retaining the specific status of maculosa have
been presented.
Although Phillips had vacillated earlier between accepting and reject-
ing the Mottled Duck and Florida Duck as distinct races, in 1924 he
synonymized the two forms. Later, Peters (1931) accepted •naculosa
as being distinct from Iulvigula, although Delacour and Mayr (1945)
as well as Hellmayr and Conover (1948) synonymized them. The form
•J, aculosa is still accepted by the A.O.U. Check-list.
Regardless of the question of the validity of the described plumage
differences, which will be discussed later, a definite hiatus in the ranges
of the populations seems to exist. The range of the Florida Duck (see
Figure 2, which is modified from Howell, 1932, and Sprunt, 1954) is
largely confined to the southern half of peninsular Florida, with its
northern limits being in the vicinity of Gainesville. There are appar-
ently no specimens of fulvigula or maculosa that have been taken in
Mississippi. A single specimen has been taken in Alabama (Imhof,
1958) and may represent either fulvigula or •nacuIosa. In addition,
several sight records along the coast of Alabama have been obtained for
Florida or Mottled ducks. Mottled Ducks occur commonly almost to
the eastern border of Louisiana (Oberholser, 1938). They are found
mainly in the coastal tier of parishes of Louisiana (Lowery, 1955) west
to the Texas border, and in Texas likewise follow the coastal counties
for the entire length of the state (Singleton, 1953). The range of the
Mottled Duck also extends a long distance into Mexico (Saunders, 1952,
1953) in the states of Tamaulipas and Veracruz. The range map pre-
sented in Figure 2 is a composite of information from the sources cited
above, and from additional sight and specimen records available to me.
The Florida Game and Fresh Water Fish Commission has taken a
census of the Florida Duck population annually since 1948 by the aerial
transect method. During the years 1948 to 1955, the estimated total
adult population varied from 17,000 to 30,000 (Sincock, 1957). The
1956 and 1957 inventory estimates for the entire state were 9,000-16,000
and 7,000-10,000 birds, respectively, indicating, according to Sincock, a
sharp drop in Florida Duck numbers in recent years. A reasonable esti-
mate of the present total adult Florida Duck population would appear to
be 10,000.
In Louisiana, inventories of the Mottled Duck population have been
made yearly since 1952. Estimates in recent years have averaged about
10,000 birds (M. M. Smith, in litt.), which are spread out fairly evenly
over the marshes of southeast and southwest Louisiana, being especially
Published in the Auk (1961) 78. Copyright 1961, University of California. Used by permission.
http://elibrary.unm.edu/sora

Citations
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Journal ArticleDOI
TL;DR: Nonindigenous species can bring about a form of extinction of native flora and fauna by hybridization and introgression either through purposeful introduction by humans or through habitat modification, bringing previously isolated species into contact.
Abstract: ▪ Abstract Nonindigenous species can bring about a form of extinction of native flora and fauna by hybridization and introgression either through purposeful introduction by humans or through habitat modification, bringing previously isolated species into contact. These phenomena can be especially problematic for rare species coming into contact with more abundant ones. Increased use of molecular techniques focuses attention on the extent of this underappreciated problem that is not always apparent from morphological observations alone. Some degree of gene flow is a normal, evolutionarily constructive process, and all constellations of genes and genotypes cannot be preserved. However, hybridization with or without introgression may, nevertheless, threaten a rare species' existence.

2,190 citations

Journal ArticleDOI
01 Oct 1986-The Auk
TL;DR: A phylogenetic analysis of all Recent genera of the Anseriformes using 120 morphological characters supports much of the current consensus regarding intraordinal relationships.
Abstract: --A phylogenetic analysis of all Recent genera of the Anseriformes using 120 morphological characters supports much of the current consensus regarding intraordinal relationships. I found that (1) Anseranas hould be placed in a monotypic family; (2) Dendrocygna, Thalassornis, geese and swans, and Stictonetta re paraphyletic to the rest of the Anatidae; (3) Cereopsis the sister group to Anser and Branta, and Coscoroba is the sister group to Cygnus and Olor; (4) Plectropterus i the sister group to the Tadorninae (shelducks) and the Anatinae (typical ducks); (5) the shelducks are monophyletic and include Sarkidiornis (provisionally), Malacorhynchus, Hymenolaimus, Merganetta, and Tachyeres; (6) the tribe \"Cairinini\" (\"perching ducks\") is an unnatural, polyphyletic assemblage and is rejected; (7) the dabbling ducks (including the smaller \"perching ducks\") comprise an unresolved, probably paraphyletic group; (8) tribal monophyly of the pochards (including Marmaronetta nd Rhodonessa), sea ducks (including the eiders), and stiff-tailed ducks (including Heteronetta) is confirmed; and (9) the retention of Mergellus and resurrection of Noraonyx are recommended based on clarifications of intratribal relationships. Problematic groups, effects of homoplasy, phenetic comparisons, life-history correlates, biogeographic patterns, and fossil species are discussed, and a phylogenetic classification of Recent genera is proposed. Received 18 November 1985, accepted 2 April 1986. THE order Anseriformes is considered to comprise the families Anhimidae (2 genera, 3 species) and Anatidae (approximately 43 genera and 150 species). The family Anatidae is undoubtedly one of the best-studied groups of birds, owing largely to the historical importance of waterfowl for hunting (Weller 1964a), domestication (Delacour 1964a), and aviculture (Delacour 1964b). The classification of the Anatidae proposed by Delacour and Mayr (1945) has been followed, with only minor revisions, in recent decades (e.g. Delacour 1954, 1956, 1959, 1964c; Johnsgard 1961a, 1962, 1965a, 1978, 1979; Woolfenden 1961; Frith 1967; Bellrose 1976; Palmer 1976; A.O.U. 1983; Bottjer 1983; Scott 1985). Perhaps the most innovative aspect of this system (inspired by the works of Salvadori 1895; Phillips 1922, 1923, 1925; and Peters 1931) was the erection of \"tribes,\" groups of genera that were considered to be closely related within the subfamilies of the Anatidae. These tribes became the primary focus of subsequent works on anatid classification, many of which addressed the tribal assignments of problematic genera (e.g. Humphrey and Butsch 1958; Johnsgard 1960a, 1961b; Humphrey and Ripley 1962; Davies and Frith 1964; Raikow 1971; Kear and Murton 1973). Most authors assumed the validity of the tribes and used them as working units in phylogenetic analyses of the family (e.g. Johnsgard 1961a, Bottjer 1983). A few workers named additional tribes (Moynihan 1958, Delacour 1959, Woolfenden 1961, Weller 1968b) or attempted to test the naturalness of those originally proposed (Cotter 1957, Woolfenden 1961, Brush 1976). Behavioral characters have been accorded considerable weight in classifications of waterfowl. Delacour and Mayr (1945) based their revision on characters they considered to be \"non-adaptive,\" including behavioral displays, nesting and feeding habits, and selected morphological characters (e.g. posture, body proportions, head shape, syringeal bulla). Reliance on comparative ethology in anatid systematics was furthered by the studies of Lorenz (19511953), McKinney (1953), and Myres (1959) and was increased significantly by Johnsgard (1960a-c, 1961a-d, 1962, 1964, 1965a, b, 1966a, b, 1967, 1978), whose work was largely ethological and influenced profoundly by that of Delacour (1954, 1956, 1959, 1964c). This emphasis, work on interspecific hybridization 737 The Auk 103: 737-754. October 1986 738 BRADLEY C. LIVEZEY [Auk, Vol. 103 (Sibley 1957; Gray 1958; Johnsgard 1960d, 1963), and study of plumage patterns of downy young (Delacour 1954, 1956, 1959; Frith 1955, 1964b; Kear 1967) were prompted in part by the opportunity to observe waterfowl in avicultural collections. Other data used in the classification of waterfowl include syringeal anatomy (Humphrey 1955, 1958; Johnsgard 1961e), cytogenetics (Yamashina 1952), serology (Cotter 1957, Bottjer 1983), osteology (DeMay 1940, Verheyen 1955, Humphrey and Butsch 1958, Woolfenden 1961, Humphrey and Ripley 1962, Raikow 1971), feather lice (Timmermann 1963), eggshell structure (Tyler 1964), egg-white proteins (Sibley 1960, Sibley and Ahlquist 1972), feather proteins (Brush 1976), royology (Zusi and Bentz 1978), lipids from the uropygial gland (Jacob and Glaser 1975, Jacob 1982), and mitochondrial DNA (Kessler and Avise 1984). These studies, with the possible exceptions of those by Lorenz (1953) and Kessler and Avise (1984), estimated the evolutionary relationships of groups by assessments of overall similarities; no attempts were made to determine primitive conditions or to distinguish shared primitive characters from shared derived characters (\"special\" similarity). Moreover, the \"evolutionary trees\" presented in most of these works lack references to the specific characters used to support the branching patterns (e.g. Delacour and Mayr 1945; Johnsgard 1961a, 1978; Woolfenden 1961). I performed a phylogenetic (cladistic) analysis of Recent genera of Anseriformes using 120 morphological characters. I present a hypothetical evolutionary tree for the order, consider the taxonomic implications, and discuss selected life-history and biogeographic correlates and the classification of selected fossil species. Many of the characters were described first in the pioneering work of Woolfenden (1961), to whom I dedicate this paper.

215 citations

Journal ArticleDOI
TL;DR: Character state mapping reconstructs monochromatic ancestors for the genus Anas as well as most of its main clades and cautions against the uncritical use of unordered parsimony as the sole criterion for inferring ancestral states.
Abstract: Although phylogenetic reconstruction of ancestral character states is becoming an increasingly common technique for studying evolution, few researchers have assessed the reliability of these reconstructions. Here I test for congruence between a phylogenetic reconstruction and a widely accepted scenario based on independent lines of evidence. I used Livezey's (1991) phylogeny to reconstruct ancestral states of plumage dichromatism in dabbling ducks (Anatini). Character state mapping reconstructs monochromatic ancestors for the genus Anas as well as most of its main clades. This reconstruction differs strongly from the widely accepted scenario of speciation and plumage evolution in the group (e.g., Delacour and Mayr 1945; Sibley 1957). This incongruence may occur because two standard assumptions of character state reconstruction are probably not met in this case. Violating either of these two assumptions would be a source of error sufficient to create misleading reconstructions. The first assumption that probably does not apply to ducks is that terminal taxa, in this case species, are monophyletic. Many of the widespread dichromatic species of ducks may be paraphyletic and ancestral to isolated monochromatic species. Three lines of evidence support this scenario: population-level phylogenies, biogeography, and vestigial plumage patterns. The second assumption that probably does not apply to duck plumage color is that gains and losses of character states are equally likely. Four lines of evidence suggest that dichromatic plumage might be lost more easily than gained: weak female preferences for bright male plumage, biases toward the loss of sexually dichromatic characters, biases toward the loss of complex characters, and repeated loss of dichromatism in other groups of birds. These seven lines of evidence support the accepted scenario that widespread dichromatic species repeatedly budded off isolated monochromatic species. Drift and genetic biases probably caused the easy loss of dichromatism in ducks and other birds during peripatric speciation. In order to recover the accepted scenario using Livezey's tree, losses of dichromatism must be five times more likely than gains. The results of this study caution against the uncritical use of unordered parsimony as the sole criterion for inferring ancestral states. Detailed population-level sampling is needed and altered transfor- mation weighting may be warranted in ducks and in many other groups and character types with similar attributes.

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183 citations

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TL;DR: Overall, the mtDNA data indicate an extremely close evolutionary relationship between Mallards and Black Ducks, and in conjunction with the geographic distributions suggest that the Black Duck is a recent evolutionary derivative of a more broadly distributed Mallard‐Black ancestor.
Abstract: We assayed restriction site differences in mitochondrial DNA (mtDNA) within and among allopatric populations of the Mallard (Anas platyrhynchos) and the American Black Duck (A. rubripes). The observed mtDNA clones grouped into two phylogenetically distinct arrays that we estimate differ by about 0.8% in nucleotide sequence. Genotypes in one clonal array were present in both species, while genotypes in the other array were seen only in Mallards. In terms of the mtDNA "gene tree," the assayed Mallards exhibit a paraphyletic relationship with respect to Black Ducks, meaning that genealogical separations among some extant haplotypes in the Mallard predate the species separation. Evidence is advanced that this pattern probably resulted from demographically based processes of lineage sorting, rather than recent, secondary introgressive hybridization. However, haplotype frequencies were most similar among conspecific populations, so the Mallard and Black Ducks cluster separately in terms of a population phenogram. The results provide a clear example of the distinction between a gene tree and a population tree, and of the distinction between data analyses that view individuals versus populations as operational taxonomic units (OTUs). Overall, the mtDNA data indicate an extremely close evolutionary relationship between Mallards and Black Ducks, and in conjunction with the geographic distributions suggest that the Black Duck is a recent evolutionary derivative of a more broadly distributed Mallard-Black ancestor.

178 citations

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