First published online September 9, 2005
Journal of Experimental Biology 208, 3581-3591 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01776
Allometric scaling of flight energetics in Panamanian orchid bees: a comparative phylogenetic approach
Charles-A. Darveau1,
Peter W. Hochachka1,
,*,
Kenneth C. Welch, Jr2,
David W. Roubik3 and
Raul K. Suarez2
1 Department of Zoology, University of British Columbia, Vancouver, BC,
Canada, V6T 1Z4
2 Department of Ecology, Evolution and Marine Biology, University of
California Santa Barbara, Santa Barbara, CA 93106-9610, USA
3 Smithsonian Tropical Research Institute, Balboa, Republic of
Panama

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Fig. 1. Phylogenetic tree hypothesized for 36 orchid bee (Apidae: Euglossini)
species, based on cyt b partial sequences and inferred using the
maximum likelihood method. Node bootstrap support values greater than 50% are
shown. Species groupings, based on Cameron
(2004 ), are presented in
different colors. Eg., Euglossa; Ef.,
Eufriesea; Ex., Exaerete; El.,
Eulaema.
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Fig. 2. Phylogenetic relationships among 15 of 18 species of orchid bees used for
hovering flight measurements. The character values are presented for body mass
Mb, wingbeat frequency n, mass-specific metabolic
rate
*CO2.
Sample size, N, is identified for each species in parentheses.
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Fig. 3. Relationships between body mass Mb and wing
morphological characters. (A) Forewing length R, (B) total wing area
S and (C) calculated wing loading Pw. Solid circles
represent the genus Euglossa; open circles, Exaerete; filled
squares, Eulaema; open squares, Eufriesea.
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Fig. 4. Relationship between body mass and hovering flight wingbeat frequency.
Symbols as in Fig. 3.
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Fig. 5. (A,C,E) Relationship between body mass, wingbeat frequency and
mass-specific metabolic rate independent contrasts obtained from cyt
b phylogeny (Fig. 1), and
using gradual (filled circles, solid lines) and speciational (open circles,
broken lines) models of character evolution. (B,D,F) These relationships
(solid and broken lines) are superimposed on the distribution of correlation
coefficients resulting from analyses performed with 10 000 different trees
(see Materials and methods).
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Fig. 6. Relationships between body mass and hovering flight whole-animal (A) and
mass-specific (B) metabolic rate. Symbols as in
Fig. 3.
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Fig. 7. Correlations between wing loading and wingbeat frequency residuals obtained
from (A) the body mass regression (r2=0.86,
F1,16=8.74, P=0.0088) and (B) from independent
contrasts for gradual (filled circles, r2=0.86,
F1,9=62.88, P< 0.0001) and speciational (open
circles, r2=0.90, F1,9=90.94,
P<0.0001) models of character evolution. Symbols in A as in
Fig. 3.
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Fig. 8. Correlations between hovering flight wingbeat frequency and mass-specific
metabolic rate residuals obtained from (A) the body mass regression
(r2=0.46, F1,16=14.25,
P=0.0015) and (B) from independent contrasts for gradual (filled
circles, r2=0.47, F1,9=8.73,
P=0.0144) and speciational (open circles,
r2=0.70, F1,9=23.73,
P=0.0007) models of character evolution. Symbols in A as in
Fig. 3.
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Fig. 9. Correlations between wing loading and mass-specific metabolic rate
residuals obtained from (A) the body mass regression
(r2=0.34, F1,16=8.23,
P=0.0110) and (B) from independent contrasts for gradual (filled
circles, r2=0.50, F1,9=9.89,
P=0.0104) and speciational (open circles,
r2=0.66, F1,9=19.11,
P=0.0014) models of character evolution. Symbols in A as in
Fig. 3.
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Fig. 10. Relationships between body mass and wing area (A), wing loading (B) and
wingbeat frequency (C) in insects ranging from fruit flies to moths (open
circles; Byrne et al., 1988 )
and orchid bees (filled circles; this study). (D) The correlation between wing
loading and wingbeat frequency residuals obtained from the body mass
regression in B and C.
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© The Company of Biologists Ltd 2005