Wilson Bull., 98(3), 1986, pp. 433444

SUBSPECIES OF THE GREATER
THEIR NAMES’RICHARDSCAUP AND

C. BANKS*ABSTRACT.-The name Fuligula mariloides Vigors, presently used by many authors for asubspecies of the Greater Scaup, Aythya marila, was originally proposed for the Lesser
Scaup, A. afinis, and may not be used in combination with the name marila. The namemariloides has been applied to a population of Greater Scaup in Kamchatka and the Commander Islands, supposedly distinguished by small size and dark dorsal color, or for that
population and the one in North America, otherwise known as A. m. nearctica. Evidence
for a subspecifically distinct population in eastern Asia is lacking, and A. marila is best
considered to consist of only the Eurasian A. m. marila and the American A. m. nearctica.
There is some interchange of the two subspecies in migration. Received 13 Dec. 1985,
accepted 4 Mar. 1986.

The American Ornithologists’ Union (AOU 1957) used the name Aythya marila nearcticaStejneger, 18 8 5, for the North American subspeciesof the Greater Scaup. More recent authorities use A. m. mariloides (Vigors, 1839) for the North American population (Bellrose 1976, Palmer
1976) or for a population in North America and eastern Asia (Johnsgard
1975, 1979). Others (Portenko 1972) recognize both an east Asian mariloides and an American nearctica in addition to the European A. m.
marih, and still others (Vaurie 1965) treat A. marila as monotypic. Whois correct?
The name Anus marila was applied by Linnaeus (176 1) to the Scaup
Duck of Europe, the species now called in North America the Greater
Scaup. In the intervening years the species has been placed in a numberof genera, but generic placement will be ignored here. Generic names
mentioned beyond, in full or as initials, are all synonymous with themodern Aythya and will be used without comment.
It was recognized early that the Scaup Duck of Linnaeus occurred also
in the New World (Pennant 1785, Wilson 18 14) and across northern Asia
(Pallas 183 1). It is only in retrospect that we are able to realize that manyof the early references to American scaup applied, totally or partly, toanother species, now known as the Lesser Scaup, A. afinis. Richardson(in Swainson and Richardson 183 1) seems to have been the first to realizethat 2 kinds of scaup occur in North America. Richardson wrote (p. 453):“Our specimens are smaller than English ones killed in the winter, the’ This paper is dedicatedto John W. Aldrich on the occasion of his 80th birthday.
2 Fish and Wildlife Service, National Museum of Natural History, Washington, D.C. 20560.

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head, bill, wings, and legs, in particular, being proportionally smaller, andthe bill less high at the base . . . but an attentive examination of a numberof specimens disclosed no peculiarities which could characterize it as a
distinct species, except its size. The undulated markings on the back andwings are darker and less extended than in the English specimens.” In
what amounts to a parenthetical statement, he also noted that “A variety,nearly corresponding with the English one in size, is also found in the
fur-countries. . . .” Wing lengths given equate to 196 mm for the smaller
birds, and 224 mm for both the larger American variety and the English
birds. In describing the bill of the scaup, Richardson (op. cit.) referred tothe unguis (nail) as “small and distinct” as in the Pochard, A. ferina, andthe Canvasback, A. valisineria. This description indicates certainly that
his basic account is about what later would be known as A. afinis.
Eyton (1838) distinguished the present Aythya a&is as an American
species distinct from marila, which he apparently considered strictly an
Old World species. Smaller size and the narrower nail on the bill formed
the main bases of the separation, as these characters still do. The type,from “North America,” is in the British Museum (Natural History)(BMNH) (Salvadori 1895, Warren 1966). One wonders if this is a birdthat Richardson had studied; the name certainly applies to the birds hehad discussed.
Vigors (1839) was the next to provide a name for a scaup population,although the name was used provisionally and the precise origin of thebirds was not stated. In reporting on the birds collected during the voyage
ofthe Blossom,under Captain F. W. Beechey, in 1825-1828, Vigors wrote
as follows under the heading Fuligula Marila: “Several specimens of abird nearly allied to this species, if not the same, were brought home by
the expedition. They uniformly differ from the typical Ful. Marila in theirsmaller size, in the black colour on the breast being less intense anddefined, in the undulating white markings being less diffused over thescapulars and back, and being wanting almost entirely on the wing-coverts. Dr. Richardson, whose judgement on these points, and whose experience respecting the birds of the Arctic region entitle him to every
confidence, is inclined to consider these birds but as a variety of theEuropean species. Following his opinion, I refrain from describing themas separate. It is however to be observed, that the true Fuligula Marila
is found in North America, and there is therefore less reason to believe
the birds alluded to above to be varieties resulting from climate or locality.
Should the species prove to be distinct, the specific name of Mariloi’des,which has been suggested by Dr. Richardson, would be appropriate.”
Publication of the results of the Beechey expedition had been delayedconsiderably, and Richardson (and perhaps others) had access to theBunksl

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specimens well before Vigors’s account appeared. Richardson was offered
the opportunity to incorporate into his own work a list of species fromthe northwestern American coast, but declined because (Swainson andRichardson 183 1:xiii) “very few of the specimens brought home on thatExpedition had notes attached to them to indicate their locality, so thatthe native places of many are uncertain.” It is obvious, however, that thesmall birds that Vigors had were the same as the smaller scaup that
Swainson and Richardson (183 1) discussed. Yarrell(1843)
discussed two
species of scaup, and used the name mariloides for the smaller American
birds without mentioning the name afinis. Yarrell both described andfigured a single specimen of what he thought was the smaller American
scaup. That bird, which had been bought in a market in England, waslater identified as a hybrid A. ferina x A. marila (Salvadori 1895). Boththe description and illustration apply remarkably well to the Redhead, A.
americana. Interestingly, Audubon (1844) gave Yarrell credit for the firstgood description of mariloides.
Eyton’s (1838) name for the small American scaup, afinis, was routinely overlooked or ignored by British authors (e.g., MacGillivray 18 52).
However, Baird (in Baird, Cassin, and Lawrence 1858) recognized affinisas the name of the smaller species, and treated mariloides Vigors as asynonym. Schlegel(l866); Gray (187 1); and Baird, Brewer, and Ridgway(1884) treated the two names in the same way. This synonymy would
seem to have been firmly established.
Meanwhile, Swinhoe (1873) detected two size classes of scaup in China,
The abundant, and apparently regularly occurring, form he recognized as
marila. Some smaller birds brought to him, from one or two flocks, poseda problem in identification. Swinhoe first considered them to be afinis,
because of their size and after consultation of works by Yarrell and Baird(presumably those cited above). Those birds, however, had extensive
white wing markings that MacGillivray(1852) and Schlegel (1866) attributed to marila rather than to a&is, and the combination of size andwing characters led Swinhoe to reject either name as applicable. He stated:“Our bird, then, is not the American F. afinis; but it nevertheless must
be the bird that Mr. Yarrell mentions under the head A‘ merican Scaup,’
which was brought home by Capt. Beechey from Behring’s Straits, andwas apparently identified with that species. Sir John Richardson suggests
for this Pacific species the name mariloides, which is generally regarded
as a synonym of aBnis. But I think it can now be shown that F. mariloides
is a species of itself, and a visitant to the Chinese coast. Fuligula marila
also occurs here in abundance. . . .” This is the first suggestion in theliterature that the birds from Beechey’s voyage were from the Bering
Straits, or that the name mariloides referred to a Pacific species. These436

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concepts may have been derived from the long title of the work in which
Vigors (1839) proposed the name mariloides, but were counter to the thengenerally accepted notion that mariloides was an American form.
In a report on birds from the Commander Islands, Stejneger (1885)
continued to associate mariloides with afinis, giving the former subspecific rank and assigning to it a range on the “Pacific coast of Asia, fromJapan southward,” where no scaup was known as a breeding bird. At thesame time, Stejneger distinguished the North American population ofGreater Scaup from the European birds, under the new name A. m. nearctica. Stejneger’s account is confusingly written, but it is clear that heconsidered his specimens from Bering Island (one of the Commander
Islands) to be A. m. marilu, and he presented measurements only for
those birds, not for the American form that he named. It seems obviousthat he took his concept of the range of mariloides from Swinhoe (1873),
not recognizing that the latter’s discussion referred only to wintering birds.
Seebohm (1890) wrote of marilu in Japan that “Fully adult males fromJapan are precisely similar to those from the British Islands.” He pointed
out that the amount of white on the wing, used by MacGillivray (1852)
and Schlegel (1866) as a specific character and by Stejneger (1885) as asubspecific character, was actually age related. Seebohm treated “Aythya
afinis mariloides Stejneger” as a synonym of marila, which he consideredmonotypic. Thus Seebohm transferred mariloides from the synonymy ofafinis to the synonymy of marila.
Salvador-i (1895) presented synonymies for the scaup following Stejneger’s (1885) distributional scheme but including both neurctica and mariloides as part of a monotypic species marila. The A.O.U. (1899) followed
Salvadori in treating both marila and a&is as monotypic.
In a discussion of the European marila, Dresser (1903) gave specificstatus to both the American nearctica and the East Asiatic mariloides,
without comment or more detailed delineation of range. Millais (19 13)
included “Kamtschatka and the Commander Isles” in the breeding rangeof marilu, noting that “These Asiatic Scaup frequenting the two lastnamed places have been separated recently under the name of F. mariloides (Vig.).” The association of those breeding grounds with mariloides
may have resulted from a careless reading of Stejneger’s (1885) work, butamounted to a compounding of the errors of several previous authors.
Millais cast doubt on the validity of both mariloides and nearctica asrecognizable forms, but for the first time treated aflnis as a subspecies ofmarila.
Hartert (1920) finalized the association of the name mariloides with
the Commander Islands-Kamchatka breeding scaup. He noted but rejected Stejneger’s arrangement of scaup populations, stating that: “I findBanksl

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that all specimens from the Far East-i.e.
from Japan and China andfrom the Bering Island-are (with few exceptions) smaller than European
N. m. marilu, the wings of 16 adult male examples measure 207-220mm., in the European form 220-230 mm. Moreover, in nearly all cases
the black barring of the upperside is coarser, thus giving the latter a darker
appearance, which is particularly noticeable on the scapulars. . . . We must
therefore, unless we suppress this form altogether, recognize a Far-East
form breeding on Bering Island, and probably in Kamtchatka, winteringin China and Japan.” Hartert recognized three subspecies of marilaneurcticu in North America; marila in Europe and northern Asia, withindefinite Asian limits; and mariloides breeding on Bering Island, probably Kamchatka and elsewhere, wintering in eastern Asia. He noted thatthe latter population is closely allied to marila, not to afinis. Hartert’s
separation of marila from mariloides at a wing length of 220 mm isdirectly opposed to Seebohm’s (1890) and Salvadori’s (1895) analyses.
Peters (193 1) followed Hartert’s classification of the scaup, and additionally fixed the type locality of mariloides by giving it as “No type
locality = Bering Sea.” No reason for this decision was given and one
might guess that Peters, as perhaps Swinhoe (1873) earlier, was influenced
by the title of the work in which mariloides was named and also by
Hartert’s use of the name for the Bering Island population. Hartert’sclassification was also accepted by Hellmayr and Conover (1948). Dement’ev and Gladkov (1952) however, did not recognize any subspecies
of mark, noting that the amount of variability in the supposedly differentiating characters was great. The A.O.U. (1957) again accepted the namenearctica for the North American population of Greater Scaup, followingPeters (193 1) and Hellmayr and Conover (1948).
Parkes (1958), noting that several authors had doubted the valid distinction of nearctica, examined the series of specimens in the American
Museum of Natural History (AMNH) “and found that nearctica is readily
recognizable by the coarse black barring of the upper parts, exactly as
characterized by Hellmayr and Conover. . . .” He considered mariloides“of eastern Asia” to be intermediate in color but smaller than either marilu
or nearctica, citing Hartert’s measurements.
Delacour (1959) recognized two subspecies of A. marila, the nominate
form in Europe and western Asia and mariloides in eastern Asia andNorth America, merging nearctica into the latter. His measurements ofwing length give mariloides a greater range of variation than marila, andone that completely encompasses the latter. Palmer (1976) followed Hellmayr and Conover (1948) and Parkes (1958) in separating American from
Eurasian birds on the basis of back color, but recognized only two subspecies. To the American population, however, he applied the name mar-

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iloides, treating nearctica as a synonym. Thus he followed Delacour inthe use of the name for the American population but not in the range
allotted to it, dropping the Asian population whose birds were supposedly
the basis of the name. Bellrose (1976) used the same names in the same
fashion. Johnsgard (1975, 1979) followed the taxonomic arrangement ofDelacour.
DISCUSSION

Two problems are revealed by the preceding taxonomic history. The
first involves the allotment of the name F. mariloides Vigors, 1839 toeither marila or afinis. The second is whether there is a recognizable
population of scaup in the Commander Islands and Kamchatka.
It is quite clear that the small birds with the narrow unguis that Richardson discussed in 183 1 and that Vigors named mariloides in 1839 were
the same, and that Eyton named that form afinis in 1838, while Vigors’s
manuscript on the voyage of the Blossom was awaiting publication. Both
European and American authors accepted mariloides as the smaller ofthe American species of scaup, often treating it as a synonym of a&is,
into the 1870s and beyond. It was not until Swinhoe (1873) confused by
Yarrell’s (1843) description of a hybrid as mariloides and other misinformation, associated that name with birds “from Behring’s Straits” thatthe concept of mariloides as an Asian population originated. This concept
was repeated by Stejneger (1885), who nonetheless maintained the association of mariloides with afinis. Once mariloides was accepted as
referring to an Asian population, it was logical to consider that the namewas synonymous with marila, because a&is
does not occur in Asia.
Thus Seebohm (1890) and Salvadori (1895), even as they denied thevalidity of the taxon mariloides, associated the name with European marila. By 1920, the “fact” that the name mariloides applied to east Siberian
birds, specifically those of the Commander Islands and nearby Ramchatka, was accepted, the only question then relating to the distinctnessof the population. That “fact” was finally given some basis when Peters(193 1) formally designated “Bering Sea” as the type locality of mariloides.
Although no locality was given for the scaup specimens that Vigors
discussed, and no type was (or has been subsequently) designated, thespecimens almost certainly originated in North America. Vigors (1839)
wrote about 102 species of birds in his report on Capt. Beechey’s voyage;
68 of these are American species, whereas only 3 are strictly Palearctic.
Of the other ducks discussed, 5 are strictly American and 4 are Holarctic
in distribution. The form of Anas crecca discussed was carolinensis, andthe only wigeon mentioned was Anas americana. At least one duck, theBufflehead, Bucephala albeola, “was found at San Francisco” and several

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439

of the shorebird specimens also were taken there. Probably most of thewaterbirds, including the scaup, were taken from large wintering flocks
in the San Francisco Bay area or in Monterey Bay, California, as theBlossom spent parts of 2 winters in that vicinity (Rosewater 1968). I
suggest San Francisco Bay, California, as the corrected type locality for
Fuligula mariloides Vigors, 1839, contra Peters (193 1).
The second problem is whether the east Asian and North American
populations of marila are distinct from the European population and, if
so, what the limits of their ranges are. The present consensus is that North
American birds differ from European birds by having coarser and more
extensive black vermiculation on the mantle (Hartert 1920, Hellmayr andConover 1948, Parkes 1958, Delacour 1959, Portenko 1972, Palmer 1976),
yielding a darker back overall. This character is restricted to adult males,a fact not always stated; no one has ever suggested that other sex or age
classes could be distinguished. My examination of specimens supports
this consensus, and the validity of nearctica, although the character does
not provide 100% separation. A small proportion of American birds arenearly as white on the mantle as European birds, as noted by Bishop(1895).
The supposed characteristics of the eastern Asian population have been
confusingly set forth by various authors, partly because of the confusion
over the specific application of the name mariloides and partly becausethat population was compared to different populations at different times.
Smaller size and darker dorsal color relative to European marila havebeen invoked as distinguishing features, as has paler color relative to
American birds.
Swinhoe (1873) first reported small scaup from Asia, but gave no measurements or any basis for his characterization of the birds. Both Seebohm(1890) and Salvadori (1895) denied that Asian birds (including those sent
to England by Swinhoe) were consistently smaller than European ones.
Hartert (1920) repeated that Asian birds were smaller “with few exceptions,” and provided ranges of measurements to support his statement.
His figures divided European and Asian birds at a wing length (adult male)of 220 mm, not allowing for any of the “exceptions.” Portenko (1972)
suggestedan average size difference and stated that “specimens with much
shorter wings predominate” among Asian birds relative to those of Europe, although the ranges of measurements given virtually overlap. No
other authors have given measurements specifically for the Asian population, except Stejneger (1885) for 2 males.
Available measurements from many sources (Fig. 1) do not indicatethat Asian birds have shorter wings. Hartert’s Far East sample is at thelower end of the range of variation, but overlaps most samples of European

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birds and, as noted above, apparently omits any exceptional birds. Portenko’s East Asian sample is small in wing length but so is his West Asian
sample compared to others, and the means of his two samples differ only
slightly. The mixed samples that include Asian birds, those of Delacour,
Palmer, and Dement’ev and Gladkov, do not, except for the last, show
any great tendency to include small birds. My own measurements of thechord of several samples (Table 1) (Fig. 1) show primarily that most otherwriters must have measured the flattened wing, and secondarily that there
is no great amount of geographic variation. I conclude that there is noevidence indicating that eastern Asian rnarila are recognizably smaller
than European or North American ones.
As indicated above, adult males of the European and North American
populations can be distinguished by back color. Hartert (1920) considered
East Asian birds to be darker than European ones, with those of America
the same or even darker. Parkes (1958) and Portenko (1972) consideredthem to be intermediate. Delacour believed Asian birds to be the sameas American ones, a view endorsed by Alexander Wetmore on examination of specimens in the British Museum (Natural History) (in litt. to
L. L. Short, FWS files).
I compared 10 adult males from Siberia, Japan, and Bering Island in
the AMNH and U.S. National Museum of Natural History (USNM) withlarger series of European and North American birds. As a series the Asianbirds seem to me to be more similar in dorsal appearance to European
specimens than to American birds, although one winter bird from Japanand 2 worn breeding birds from Bering Island might be classed as intermediate. I also examined several first-year males from Asia and notedthat they tend to be darker than the adults, approaching the American
birds. It is possible that age classes were not adequately separated in someprevious comparisons. I conclude that Asian birds cannot be distinguishedfrom European ones on the basis of dorsal color any more than they canon size. I recommend, therefore, that only two subspecies of Greater Scaup
be recognized, A. m. marila of Eurasia and A. m. nearctica of North
America.
DISTRIBUTIONAL

RECORDS

The Pribilof Islands represent a landfall for many migrant birds fromboth North America and Asia, and have provided the first and only
American records of several Old World species or subspecies. I haveexamined 5 adult males ofAythya marila from the Pribilof Islands in the
USNM and the University of Kansas Museum of Natural History. One
specimen taken in late April and 3 in early June are dark dorsally liketypical North American birds. One specimen, USNM 496822, taken 29

Bunksl

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441

SCAUP

AREA

N

SOURCE

Nearctic

12

I

Nearctic-2

Nearct ic
a E. Asia-

3

Bering Island

2

Far East
E. Asia

4

1

5

111

6

I

Europe61 Japan

13

W. Asia

I3

Europe45

Europe65aEurope-3Europe-9Europe

-

5

All

-

IO

All

51

I
6

I

7

II

I
8

I

200

210
WING

220

230

LENGTH(mm)

240

FIG. 1. Measurements of wing length of male (presumably adult, but not stated in every
instance) Greater Scaup. Sources: 1, Palmer (1976); 2, Oberholser (1974); 3, Delacour ( 1959);
4, Stejneger (1885); 5, Hartert (1920); 6, Portenko (1972); 7, Cramp and Simmons (1977);
8, Bauer and Glutz von Blotzheim (1969); 9, Witherby et al. (1939); 10, Dement’ev andGladkov (1952); 11, this study, Table 1. A dash under N indicates an unspecified sample
size.

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TABLE 1
WINGCHORD(MM)OFADULTMALESOFPOPULATIONSOFGREATERSCAUP,MEASUREDBYTHEAUTHOR
Area

N

Range

Meall

North America
EuropeEasternAsia
Pribilof Islands
All areas

25
12
10
4
51

205.5-222.5
206.5-218.0
205.0-223.5
211.0-217.0
205.5-223.5

215.7
214.6
210.9
214.6
213.9

June 1968 on St. George Island by Max C. Thompson, is much paler on
the back and is identical in that feature to birds from Europe and Japan.
I consider this bird a representative of the Eurasian population, A. m.
marila. Despite the date, this bird had small testes, and the species apparently does not breed on the Pribilofs. This appears to be the only
record of the Eurasian subspecies in North America. The pale color ofthis individual, incidentally, supports the rejection of the concept of adarker east Asian subspecies; it is unlikely that a European bird would
have reached the Pribilofs.
Greater Scaup from North America occasionally migrate, or wander,
to Asia. One such bird was banded on 17 September 1977 on Amchitka
Island in the Aleutians and recovered on 21 October 1977 on the islandof Hokkaido, Japan. This bird was a female that was banded in the year
it hatched and probably was a locally hatched bird although it was capableof flight and may have reached Amchitka from another area (C. F. Zeillemaker, pers. comm.). This is the only record ofA. m. nearctica in Japanor elsewhere in Asia.
ACKNOWLEDGMENTS
M. B. Hays prepared Fig. 1 and helped in the preparation of this report in many ways.
K. Klimkiewicz of the Bird Banding Laboratory of the Fish and Wildlife Service provided
the initial information on the banding recovery in Japan, and C. F. Zeillemaker of the
Aleutian IslandsUnit of the Alaska Maritime National Wildlife Refugeprovided the detailed
information and permission to report it here. J. W. Aldrich and D. W. Steadman read themanuscript in an early form and offered helpful suggestions.Authorities at the American
Museum of Natural History, the Academy of Natural Sciencesof Philadelphia, and theMuseum of Natural History at the University of Kansas allowed me to study birds in theircollections.I particularly appreciate the time and effort spent by I. C. J. Galbraith of the
British Museum (Natural History) in attempting, unsuccessfully,to locate a specimenthat
might be the type of Fuligula mariloidesVigors.

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LITERATURE

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443

CITED

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AUDUBON, J. J. 1844. The birds of America. Vol. 7. J. J. Audubon, Philadelphia, Pennsylvania.
BAIRD, S. F., T. M. BREWER,AND R. RIDGWAY. 1884. The water birds of North America.
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J. CASSIN, AND G. N. LAWRENCE. 1858. General report upon the zoology of theseveral Pacific railroad routes. Birds. Reports of explorations and surveys. Vol. 9, Pt.
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DELACOUR, J. 1959. The waterfowl of the world. Vol. 3. Country Life Ltd., London,
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Israel.)DRESSER,H. E. 1903. A manual of Palearctic birds. Pt. 2. H. E. Dresser, London, England.
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GRAY, G. R. 187 1. Hand-list of genera and species of birds. Pt. 3. British Museum (Nat.
Hist.), London, England.
HARTERT, E. 1920. The birds of the Commander Islands. Novitates Zoologicae 27:128158.
HELLMAYR, C. E. AND B. CONOVER. 1948. Catalogue of birds of the Americas and theadjacent islands. Field Mus. Nat. Hist., Zool. Ser., Vol. 13, Pt. 1, No. 2.
JOHNSGARD,P. A. 1975. Waterfowl of North America. Indiana Univ. Press, Bloomington,
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1979. Order Anseriformes in Check-list of birds of the world. Vol. 1, 2nd ed. (E.
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MACGILLIVRAY, W. 1852. A history of British birds. Vol. 5. William S. Orr and Co.,
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PENNANT, T. 1785. Arctic zoology. Vol. 2. Class 2. Birds. Henry Hughs, London, England.
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