The Wilson Journal of Ornithology 123( 1 ):76—84, 2011 
GEOGRAPHIC SONG VARIATION IN THE NON-OSCINE CUBAN 
TODY (TOD US MULTICOLOR) 
ENEIDER E. PEREZ MENA'- 3 AND EMANUEL C. MORA 2 
ABSTRACT.—We studied sound emission in the non-oscine Cuban Tody (Todus multicolor) to quantify its acoustic 
repertoire and to document geographic variation in its songs across the Cuban archipelago. Cuban Todies emitted three 
types of sounds. The characteristic song of the species was recorded from 98% of 116 individuals. The characteristic song 
of the species and a variant form recorded from two individuals consisted of trains of multi-harmonic short, downward 
frequency modulated notes emitted at peak frequencies below 4 kHz. A third type of sound in the limited repertoire of the 
species recorded from two birds is presumably produced with the wings and appears in the spectrograms as a train of short 
clicks with frequencies also below 4 kHz. Evidence of geographic variation was found in the characteristic song. Birds from 
Isla de la Juventud and Pinar del Rio emitted more notes per train spaced at longer intervals than birds from the rest of the 
provinces. The peak frequency of the notes had lower values in birds from Isla de la Juventud. A discriminant function 
analysis grouped todies from different provinces into two main clusters corresponding to western Cuba and eastern Cuba. 
This geographic song variation may indicate genetic differences in this sedentary forest bird, and the existence of two 
incipient species of todies in Cuba. Isolation may have been caused by discontinuities in the mainland of Cuba that 
occurred between the Pleistocene and Holocene or by deforestation occurring in Cuba for the last five centuries. Received5 
January 2010. Accepted 19 October 2010. 
Geographical variation should be considered 
when analyzing vocal repertoires of birds, as 
species distributed across wide areas may vary in 
their songs. Geographic variations have been 
reported mostly from oscines (Mundinger 1982, 
Martens 1996), but also from non-oscine (Gold¬ 
stein 1978, Saunders 1983, Bretagnolle 1989, 
Wright 1996) and suboscine taxa (Isler et al. 1998, 
Lindell 1998, Isler et al. 1999). 
Several theories have been proposed to explain 
the geographic variation in vocalizations observed 
in different species. Geographic variations may 
result lrom adaptation to local conditions, such as 
social or structural environments (Lemon 1975, 
Payne 1980), but also from non-adaptive process¬ 
es such as the accumulation of genetic or cultural 
mutations in a population due to isolation or 
founder events (Tack et al. 2005), or some 
combination of genetic variation and learning 
(Kroodsma 1996). Geographic variation in vocal¬ 
izations is expected to be reinforced by limited 
intermixing between the species’ populations due 
to geographic isolation. Thus, geographic son* 
variation may reflect structure among populations 
of a species. 
The Cuban Tody (Todus multicolor) belongs to 
'Institute for Ecology and Systematic, Carretera de 
Varona Km 3 I/2 , Capdevila, Boyeros, CP 10800, Ciudad de 
La Habana, Cuba. 
Faculty of Biology, Havana University, Calle 25, No 
455 entre J e I. Vedado, Plaza de La Revolution, CP 10400 
Ciudad de La Habana, Cuba. 
Corresponding author; e-mail: jrrubio@infomed.sld.cu 
a genus of charismatic small and colorful non- 
oscine birds endemic to the Caribbean, represent¬ 
ed by two species in Hispaniola, one in Jamaica, 
one in Puerto Rico, and one in Cuba (Raffaele et 
al. 1998). The Cuban Tody is well distributed in 
Cuba and is common in semi-deciduous meso- 
phytic forest and coastal vegetation (Garrido and 
Kirkconnell 2000), swamp vegetation complex 
(Gonzalez et al. 1997), and secondary vegetation 
(Gonzalez et al. 2001). This species perches for 
long periods, often bobbing its head up and down 
while locating the small adult and larval insects, 
and spiders upon which it feeds. The sedentary 
behavior, forest use, and short distance flight of 
this species suggest easier isolation of populations 
by deforestation, a phenomenon that has occurred 
in Cuba during the last five centuries (Fig. 1)- ft 
may be advantageous to study the song repertoire 
of the Cuban Tody since, among many non- 
oscines, songs are not learned from other 
individuals (Konishi and Nottebohm 1969, 
Kroodsma 1989), and vocal behavior may be 
used as an unambiguous genetic marker for an 
individual. 
Cuba, the largest archipelago of the Greater 
Antilles, is <150 km wide but extends east-west 
lor >1,000 km. We tested the hypothesis that the 
song of the Cuban Tody exhibits geographic 
variation across the length of Cuba by character¬ 
izing the acoustic repertoire of the species from 
individuals recorded in seven provinces represen¬ 
tative of the island. We specifically predicted that 
song similarity should be inversely related to 
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