Bc. Livezey, AN ECOMORPHOLOGICAL REVIEW OF THE DODO (RAPHUS-CUCULLATUS) AND SOLITAIRE (PEZOPHAPS-SOLITARIA), FLIGHTLESS COLUMBIFORMES OF THE MASCARENE ISLANDS, Journal of zoology, 230, 1993, pp. 247-292
This paper describes a morphological study of the dodo Raphus cucullat
us and solitaire Pezophaps solitaria-extinct, flightless Columbiformes
of the Mascarene Islands, Indian Ocean--based on mensural data from 3
87 skeletal elements, comparative data from four flighted species of C
olumbidae, ancillary mensural data from other Columbiformes, and the l
iterature. Raphus cucullatus and P. solitaria are characterized by gre
at body size and substantially reduced pectoral limbs. Sexual size dim
orphism is unusually great in both R. cucullatus and P. solitaria, and
sexual dimorphism of P. solitaria may be the greatest of any carinate
bird. Estimates of body mass (kg), based on femur lengths of flighted
columbids and adjusted for flightlessness and seasonal deposition of
fat, were: 21 and 17 for male and female R. cucullatus. and 28 and 17
for male and female P. solitaria, respectively. Pectoral reduction is
greater in R. cucullatus than in P. solitaria, but both species are ch
aracterized by differential shortening of wing elements, changes in th
e sternum and scapulocoracoidal angle. R. cucullatus has disproportion
ately long femora, short tarsometatarsi and long digits, whereas P. so
litaria has disproportionately long tibiotarsi, short tarsometatarsi a
nd short digits; relative shaft widths of leg elements are substantial
ly greater in both R. cucullatus and P. solitaria than in flighted col
umbids. As is typical of other birds, giantism of R. cucullatus and P.
solitaria probably was associated with physiological changes, increas
ed longevity, enhanced thermodynamic efficiency and improved capacity
for fasting. Evidently both R. cucullatus and P. solitaria were primar
ily frugivorous, and foraging-related morphological peculiarities incl
ude an enlarged crop and the retention of 'gizzard stones'. Both speci
es had clutch sizes of one, and egg masses of the flightless species a
pproximated that predicted for a flighted columbid of equal size. Extr
eme sexual dimorphism and territoriality of P. solitaria suggest that
the species may have been polygynous and perhaps lek-breeding. The des
cription of 'gigantic immaturity' in R. cucullatus by Strickland & Mel
ville (1848) probably represents the first recognition of paedomorphos
is in any species of bird. Pectoral underdevelopment and (in at least
R. Cucullatus) comparatively 'juvenile' plumage in adults substantiate
the role of paedomorphosis in the ontogeny of flightlessness in R. cu
cullatus and P. solitaria; both species also were characterized by per
amorphic skulls, trunks and pelvic appendages. The common assumption o
f monophyly of the flightless species (i.e. Raphidae) lacks rigorous a
nalytical support. In spite of the anthropogenous extinction of both s
pecies, both R. cucullatus and P. solitaria were evolutionarily innova
tive in ontogeny, morphological characters and life-history strategies
.