THE anterior margin of the orbit in ducks is formed by the lacrimal bone,
which articulates with the anterolateral margin of the frontal bone and the
posterodorsal corner of the maxillary process of the nasal bone. The evolu-
tion of this bone in birds generally has been dealt with recently by
Cracraft (Amer. Midl. Naturalist, 80: 316, 1968), whose terminology for
the parts of the lacrimal bone is followed here. In many ducks the postero-
dorsal corner of the lacrimal is marked by a small tubercle, which serves
as the site of attachment of the anterior end of the orbital membrane, a
sheet of connective tissue that covers and protects the dorsal aspect of the
eyeball. In some forms this tubercle has become elongated to form a
stout, finger-like projection, the supraorbital process. This appears to
provide mechanical protection to the eyeball and salt gland (Figure 1).
Table 1 lists the occurrence and degree of development of this process
in the skulls of all living genera of ducks.
CORRELATION WITH FEEDING AND LOCOMOTOR HABITS
In the following discussion, data on feeding habits are from Delacour
(The waterfowl of the world, vols. 1-3, London, Country Life Ltd., 1954,
1956, 1959). Table 1 shows that the supraorbital process is developed
significantly only in certain groups of ducks that feed underwater. It is
absent or rudimentary in the Tadornini, Cairinini, and Anatini, which are
primarily surface feeders, but also in Merganetta arma.ta, the Torrent
Duck, which feeds underwater. In the Aythyini it is fairly well-developed
in several species of Aythya, which are excellent divers, but is rudimentary
in Netta peposaca, which is more of a surface feeder. It is also rudimentary
in the Canvasback, Aythya valisineria, which dives for vegetation. Among
the Mergini the supraorbital process is highly developed in eiders (Poly-
sticta, Somateria) , scoters ( Melanitta) , and Long-tailed Duck ( Clangula
byemalls), all of which feed mainly on invertebrates taken from the
bottom. It is also fairly large in the Harlequin Duck, Histrionicus his-
trio.nicus, which feeds on invertebrates, often in turbulent waters. In
contrast, the process is absent or rudimentary in the Bufflehead and
goldeneyes (Bucephala) and the closely related mergansers (Mergus).
These species are largely carnivorous, the mergansers eating mostly fish.
Among the stifftail ducks (Oxyurini) the supraorbital process is unde-
veloped in both the Black-headed Duck (Heteronetta atricapilla) and the
Ruddy Duck (Oxyura jamaicensis) which feed mainly on vegetation, and
SALT GLAND
Figure 1. Lacrimal bone and adjacent structures in a male King Eider, Somateria
s pectabilis.
in the carnivorous Musk Duck (Biziura lobata). In the steamer ducks
(Tachyerini) the process is quite large, resembling that in the eiders. Both
groups feed widely on marine invertebrates.
In general, the supraorbital process is most highly developed in those
forms that feed on the bottom and that might be likely to damage the
eye by abrasion against underwater objects such as the hard shells in
mollusk beds. Freshwater divers that feed on plants or invertebrates in the
soft mud of ponds or lakes are probably less subject to such injury. The
mergansers, which pursue fish in open water, are less exposed to the
possibility of such eye injuries.
EVOLUTION
The most recent studies of waterfowl evolution (e.g. Johnsgard, Water-
fowl/their biology and natural history, Lincoln, Univ. Nebraska Press,
196.8, p. 3) suggest that the Oxyurini, Mergini, Aythyini, and Tachyerini
arose independently from some Anatini-like, surface feeding ancestral
group. Hence specialization for diving and underwater food-gathering
arose at least four times in the history of the Anatinae, and the develop-
ment of an elongated supraorbital process presumably occurred three times.
It is not clear why this structure did not arise in such bottom-feeding
forms as the Oxyurini or Merganetta armata. Perhaps the necessary
genetic changes upon which selection could operate simply never took
place.
The question arises as to whether the elongated process should be con-
sidered homologous in the different groups of ducks. It is generally held
TABLE 1
OCCURRENCE AND SIZE 1 OF TIlE SUPRAORBITAL PROCESS IN DUCKS
Actual length Relative length s
Species No. Range Mean Range Mean
TADORNINI
C yanochen cyanopterus 1
Chloephaga melanoptera 2
Neochen jubatus 1
Alopochen aegyptiacus 1
Tadorna tadornoides 1
TACHYERINI
Tachyeres patachonichus 2 3.3-17.0 10.2 14-75 44
T. pteneres 1 6.3-7.0 6.7 24-25 24.5
T. brachypterus 1 8.4 8.4 - 49
CAIRININI
Plectropterus gambensis 1 Absent
Cairina moschata 2 Rudimentary
Pteronetta hartlaubl 1 Rudimentary
Sarkidiornis melanotos 1 Absent
Nettapus coromandelianus 1 Rudimentary
Callonetta leucophrys 1 Absent
Aix sponsa 4 Absent
Chenonetta jubata 2 Absent
Amazonefta braziliensis 1 Absent
ANAT1NI
Merganetta armata 1 Absent
Hymenolaimus malacorhynchus 1 Absent
Anas crecca 5 Absent or rudimentary
A. gibberifrons 3 Absent or rudimentary
A. platyrhynchos 5 Absent or rudimentary
A. rubripes 2 Rudimentary
Malacorhynchus membranaceus 3 Rudimentary
Marmaronetta anguirostris 1 Absent
AYTHYINI
Rhodonessa caryophyllacea 1 Rudimentary
Netta peposaca 2 Rudimentary
A ythya valisineria 4 Rudimentary
A. americana 5 0.7-1.8 1.3 3-9 6
A. marila 3 1.1-2.3 1.6 6-12 8
A. a]finis 5 1.3-2.7 2.1 7-14 11
MERGINI
Somateria mollisima 4 4.7-8.9 6.2 18-33 25
S. spectabilis 3 4.6-5.2 4.9 19-23 21
S. fischeri 4 6.0-8.2 6.7 25-34 28
Polysticta stelleri 5 3.0-3.6 3.3 16-19 18
Histrionicus histrionicus 3 1.8-2.7 2.2 10-14 12
Clangula hyemalis 5 2.6-4.1 3.4 13-19 16
Melanitta nigra 3 3.7-5.0 4.6 18-24 22
M. perspicillata 5 2.3-4.0 3.2 13-18 15
M. ]usca 5 2.6-4.0 3.1 12-19 15
Rudimentary 3
Absent
Rudimentary
Absent
Rudimentary
x Measurements in mm.
-øThis is the actual length divided by the width of the orbit and expressed as a per-
centage. The mean relative length is most useful for the purpose of comparing the
development of the process in different species.
a The process is considered rudimentary if less than 1.5 mm. in length.
Supraorbital Process in Ducks
TABLE 1 (Continued)
Actual length
Species No. Range Mean'
Relative length s
Rtnge Mean
Bucephala albeola 4 Absent or rudimentary
B. islandica 1 Absent
B. clangula 4 Absent
Mergus albellus 1 Rudimentary
M. serrator 5 Rudimentary
M. merganser 3 Absent or rudimentary
OXYURIXI
Heteronetta atricapilla 2 Absent or rudimentary
Oxyura jamaicensis 5 Absent or rudimentary
Biziura lobata 3 Absent or rudimentary
that homology occurs when the condition or feature is also found in a
form ancestral to the groups under discussion. As it is believed that the
different tribes of diving ducks evolved separately from a surface-feeding
ancestry, the condition would not be considered homologous by this defini-
tion alone. It seems probable that in each case this advanced condition
resulted by parallel evolution from a common ancestral structure, the
tubercle of attachment of the orbital membrane. Bock (Amer. Naturalist,
47: 265, 1963) suggests that, in closely related forms, features that appear
to be homologous may have evolved independently in groups whose
common ancestor possessed a precursor of the advanced condition. This
TABLE 2
FAMILIES AND SPECIES OF BIRDS EXAMINED IN ADDITION TO THOSE IN TABLE 1
Spheniscidae:
Gaviidae:
Podicipedidae:
Diomediidae:
Procellariidae:
Pelecanoididae:
Pelecanidae:
Sulidae:
Phalacrocoracidae:
Anhingidae:
Fregatidae:
Anseranatidae:
Anatidae:
Rallidae:
Heliornithidae:
Alcidae:
Cinclidae:
Pygoscelis adeliae; Spheniscus humboldti; S. magelIanicus; S.
mendiculus ; Megadyptes antipodes; Eudyptula minor; Aptenodytes
]orsteri.
Gayla adamsii; G. arctics; G. iraruer; G. stellata.
Aechmophorus major; A. occidentalis; Centropelma micropterum;
Podiceps auritus; P. caspicus; P. cristatus; Podilymbus podiceps.
Diomedes exulans ; D. immutabilis ; D. nigripes.
Daption capensis ; Macronectes giganteus ; Pterodroma leucopterus ;
Pachyptila ]orste.
Pelecanoides garotii.
Pelecanus conspiciIlatus.
Sula dactylatra.
Nannopterum harrisi.
Anhinga novae-hollandiae.
Fregata magnificens.
A nseranas semipalmata.
Anser albi]rons; Branta canadensis; B. bernicla; Cygnus strata;
Coscoroba coscoroba; Dendrocygna autumnalis; D. bicolor.
Fulica americana.
Heliornis ]ulica.
Pinguinus impennis ; Uria aalge ; Cerorhinca monocerata.
Cinclus mexicanus.
would indicate that the descendent groups share an evolutionary potential
inherited from the ancestor, and the advanced condition should then be
considered to be homologous. The supraorbital process in ducks appears
to be such a case.
In order to ascertain whether a similar development has occurred in
other groups of aquatic birds, I surveyed the skulls of representative species
of a wide variety of orders (Table 2). In most cases no structure exists
comparable to the elongated supraorbital process of ducks, although some
penguins and grebes have a rudimentary process that presumably functions
as a place of attachment of the orbital membrane. Albatrosses (Diomedea)
and some Procellariidae (Dap.tion, Pterodro.ma) have a short, blunt process
that may provide protection similar to that suggested for ducks. A short
process also is present in the Finfoot, Heliornis Julica. Among Anatidae
other than true ducks (Anatinae), a process of more than rudimentary size
was found only in the swanlike Coscoroba coscoroba. Its significance in
this nondiving, surface-feeding species is not known.
ACKN OWLEDGVfENTS
I would like to thank Nicolaas Verbeek and Wilbur Quay for reading and criticizing
the manuscript, and Peter Ames for discussing the problem with me. F. B. Gill
kindly measured specimens of Tachyere$ in the collection of the University of Michi-
gan Museum of Zoology. Gene M. Christman offered helpful advice in the prepara-
tion of the illustration. I am also grateful to the following persons for loaning or
providing access to specimens in their care: Dean Areadon, American Museum of
Natural History; D. W. Snow, British Museum (Natural History); Robert W. Storer,
University of Michigan Museum of Zoology, and Richard L. Zusi, U.S. National
Museum.
Museum oJ Vertebrate Zoology, University oJ CaliJornia, Berkeley,
CaliJornia 94720.