The following article is recently published and has free access in:

http://www.biomedcentral.com/1471-2148/7/136


Laura J. May-Collado, Ingi Agnarsson, Douglas Wartzok. 2007. Phylogenetic 
review of tonal sound production in whales in relation to sociality. BMC 
Evolutionary Biology 7:136.

Abstract 
Background
It is widely held that in toothed whales, high frequency tonal sounds called 
'whistles' evolved in association with 'sociality' because in delphinids they 
are 
used in a social context. Recently, whistles were hypothesized to be an 
evolutionary innovation of social dolphins (the ‘dolphin hypothesis’). However, 
both 'whistles' and 'sociality' are broad concepts each representing a 
conglomerate of characters. Many non-delphinids, whether solitary or social, 
produce tonal sounds that share most of the acoustic characteristics of 
delphinid whistles. Furthermore, hypotheses of character correlation are best 
tested in a phylogenetic context, which has hitherto not been done. Here we 
summarize data from over 300 studies on cetacean tonal sounds and social 
structure and phylogenetically test existing hypotheses on their co-evolution.
Results
Whistles are 'complex' tonal sounds of toothed whales that demark a more 
inclusive clade than the social dolphins. Whistles are also used by some 
riverine 
species that live in simple societies, and have been lost twice within the 
social 
delphinoids, all observations that are inconsistent with the dolphin hypothesis 
as stated. However, cetacean tonal sounds and sociality are intertwined: (1) 
increased tonal sound modulation significantly correlates with group size and 
social structure; (2) changes in tonal sound complexity are significantly 
concentrated on social branches. Also, duration and minimum frequency 
correlate as do group size and mean minimum frequency.
Conclusions
Studying the evolutionary correlation of broad concepts, rather than that of 
their 
component characters, is fraught with difficulty, while limits of available 
data 
restrict the detail in which component character correlations can be analyzed 
in 
this case. Our results support the hypothesis that sociality influences the 
evolution of tonal sound complexity. The level of social and whistle complexity 
are correlated, suggesting that complex tonal sounds play an important role in 
social communication. Minimum frequency is higher in species with large 
groups, and correlates negatively with duration, which may reflect the 
increased 
distances over which non-social species communicate. Our findings are 
generally stable across a range of alternative phylogenies. Our study points to 
key species where future studies would be particularly valuable for enriching 
our 
understanding of the interplay of acoustic communication and sociality.
Laura J. May-Collado

Ph.D. Candidate
Department of Biological Sciences
Florida International University
11200 SW 8th Street, Miami, FL 33199
Email: [EMAIL PROTECTED]
Personal Home Page: www.delphinids.com

Investigadora Asociada 
Fundacion KETO
Apartado 1735-1002 San Jose, Costa Rica
www.fundacionketo.org
Phone: (305) 348-7429

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