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Environmental and behavioral influences on the comparative energetics of Anhingas, Double-crested Cormorants, and Flightless Cormorants

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Environmental and behavioral influences on the comparative energetics of Anhingas, Double-crested Cormorants, and Flightless Cormorants
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Hennemann, Willard W., 1954-
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v, 95 leaves : ill., maps ; 28 cm.

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Subjects / Keywords:
Ambient temperature ( jstor )
Birds ( jstor )
Body temperature ( jstor )
Drying ( jstor )
Incident radiation ( jstor )
Insolation ( jstor )
Plumage ( jstor )
Solar radiation ( jstor )
Sun ( jstor )
Thermoregulation ( jstor )
Anhinga anhinga ( lcsh )
Cormorants ( lcsh )
Dissertations, Academic -- Zoology -- UF
Zoology thesis Ph. D
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bibliography ( marcgt )
non-fiction ( marcgt )

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Thesis:
Thesis (Ph. D.)--University of Florida, 1982.
Bibliography:
Bibliography: leaves 91-94.
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Typescript.
General Note:
Vita.
Statement of Responsibility:
by Willard W. Hennemann, III.

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ENVIRONMENTAL AND BEHAVIORAL INFLUENCES ON THE COMPARATIVE
ENERGETIC OF ANHINGAS, DOUBLE-CRESTED CORMORANTS,
AND FLIGHTLESS CORMORANTS




BY



WILLARD W. HENNEMANN, III


A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE
UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY


UNIVERSITY OF FLORIDA


1982




ENVIRONMENTAL AND BEHAVIORAL INFLUENCES ON THE COMPARATIVE
ENERGETICS OF ANHINGAS, DOUBLE-CRESTED CORMORANTS,
AND FLIGHTLESS CORMORANTS
BY
WILLARD W. HENNEMANN, III
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE
UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1982


ACKNOWLEDGEMENTS
I thank my committee members for their support and advice:
Pierce Brodkorb, John Kaufmann, J.D. Martsolf, and especially my chair-
main, Brian McNab. Their considerable investment in me and intellectual
stimulation of me shaped my maturation as a scientist.
Ralph Heath and the Suncoast Seabird Sanctuary provided me with
Double-crested Cormorants. Steve Thompson provided friendship, advice,
and served as reviewer for numerous manuscripts and letters. Mike
Konecny gave me, in addition to friendship, invaluable assistance in
many phases of the project; without his help the Galapagos segment of
the study would never have been attempted.
I am also indebted to Thomas Emmel, John Anderson, Frank Nordlie,
and Carmine Lanciani for the many favors they have done for me in the
past, and for the support and encouragement given me.
Most of all I would like to thank my wife, Jeanne, my parents, and
my family. I thank Jeanne for allowing me the freedom to work odd
hours, travel to distant places, and lead the far from "normal" life of
a biologist, while still being a constant source of love, understanding,
and constructive criticism. I thank my parents and my family for pro
viding me with an enriched environment in which to develop, financial
and emotional support, and just the right amount of direction. Without
their help my graduate career and dissertation research would not have
been possible.
This research was funded by grants from Sigma Xi, the Frank M.
Chapman Memorial Fund, and NSF (# DEB 7906056).


TABLE OF CONTENTS
ACKNOWLEDGEMENTS ii
ABSTRACT iv
CHAPTER
ONE INTRODUCTION 1
TWO METHODS AND STUDY SITES 5
Field Studies 5
Laboratory Studies 8
THREE RESULTS: FIELD OBSERVATIONS 13
Anhingas 13
Double-Crested Cormorants 18
Flightless Cormorants 23
FOUR LABORATORY RESULTS 31
Energetics: Day versus Night 31
Energetics: Incident Radiation 41
Evaporative Water Loss 54
Plumage Wetting 63
FIVE DISCUSSION FIELD OBSERVATIONS 65
SIX DISCUSSIONLABORATORY RESULTS . 71
SEVEN ENERGETICS, BEHAVIOR, AND ZOOGEOGRAPHY 77
LITERATURE CITED 91
BIOGRAPHICAL SKETCH
95


Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment
of the Requirements for the Degree of
Doctor of Philosophy
ENVIRONMENTAL AND BEHAVIORAL INFLUENCES ON THE COMPARATIVE
ENERGETICS OF ANHINGAS, DOUBLE-CRESTED CORMORANTS,
AND FLIGHTLESS CORMORANTS
By
Willard W. Hennemann, III
August 1982
Chairman: Brian McNab
Major Department: Zoology
The behavior of free-living Anhingas (Anhinga anhinga), Double-
crested Cormorants (Phalacrocorax auritus), and Flightless Cormorants
(Phalacrocorax harrisi) was monitored in the field simultaneously with
ambient temperature and solar intensity. The frequency of spread
winged behavior in A. anhinga and P_. auritus increased at higher solar
intensities and was inversely correlated with ambient temperature;
both relationships were much stronger in Anhinga. Spread-winged be
havior in P. harrisi showed no apparent relation with solar intensity
and was positively correlated with ambient temperature. Anhingas ex
hibiting spread-winged postures oriented with their backs to the sun,
maximizing the surface area exposed to insolation, and maintaining their
back normal to the sun's rays. Phalacrocorax auritus and P_. harrisi
also oriented nonrandomly, but, unlike A. anhinga, less than 50% of
the cormorants observed "sunning" oriented with their backs to the sun.
Anhinga anhinga and P_. auritus faced into the sun during guiar flutter,
iv


thereby minimizing the surface area exposed to the sun and reflecting
much of the incident radiation.
Laboratory measurements showed that A. anhinga has a lower basal
metabolic rate and a higher thermal conductance than expected from body
mass, while P_. auritus has a higher basal metabolic rate and a higher
thermal conductance than expected. Both species had lower basal
metabolic rates, body temperatures, and thermal conductances at night.
Basal metabolic rates and rates of evaporative water loss of both
species were elevated under simulated solar radiation as compared to
daytime without incident radiation. Both species have elevated meta
bolic rates and thermal conductances while wet. In the laboratory and
in the field, P_. auritus only exhibited spread-winged postures when
wet; A. anhinga readily displayed such postures regardless of plumage
condition. Thermal conductance in cormorants is unchanged during
exposure to incident radiation, but Anhingas use spread-winged pos
tures to enhance insolation absorption, thereby reducing their thermal
conductance and metabolic rates at low ambient temperatures, and
extending the thermoneutral zone. These physiological and behavioral
differences between Anhingas and Double-crested Cormorants may explain
the differences in their geographic distributions.
v


CHAPTER ONE
INTRODUCTION
Few quantitative data are available concerning thermoregulation
in anhingas (Anhingidae) and cormorants (Phalacrocoracidae), two
closely related families of the Order Pelecaniformes. Bartholomew
et al. (1968) examined guiar fluttering as a mechanism for evaporative
cooling in a series of birds including cormorants, and King and Farner
(1961) reported the body temperatures of the Great Cormorant
(Phalacrocorax carbo) and the Double-crested Cormorant (Phalacrocorax
auritus) as 39.8 degrees C and 40.1 degrees C, respectively. However,
virtually nothing has been published concerning the basal metabolic
rates, thermoneutral zones, and overall metabolic responses of these
birds to fluctuating ambient temperature.
Thermoregulation in these two families is especially interesting
because of their conspicuous spread-winged behavior. Both the Double-
crested Cormorant and Anhinga (Anhinga anhinga), common to Florida,
utilize a spread-winged posture, the function of which has been the
subject of considerable debate (see Clark 1969, Kennedy 1969 for
reviews). Spread-winged postures are found primarily within the
Ciconiiformes (Clark 1969, Curry-Lindahl 1970, Kahl 1971), Falconi
formes (Clark 1969, Hatch 1970, Cade 1973, Grier 1975) and Pelecani-
formes (Snow 1966, Clark 1969, Curry-Lindahl 1970). Several functions
have been proposed for this spread-winged posture including balancing
(Rijke 1968, Kahl 1971), feather maintenance, shading of nest contents
1


2
and use as a social display (Kahl 1971), molt and skin conditioning
(Potter and Hauser 1974), wing drying (Rijke 1968, Kennedy 1969,
Kahl 1971), vitamin D production (Kennedy 1968, 1969), ectoparasite
removal (Kennedy 1969), and thermoregulation (Heath 1962, Clark 1969,
Kennedy 1969, Kahl 1971, Siegfried et al. 1975, Berry 1976). All of
these proposed functions may apply in certain species under certain
conditions; that is, there may be no one function for this behavior.
However, the two most commonly cited functions, which have been pro
posed for the greatest number of species (including anhingas and
cormorants), are wing drying and sunning for thermoregulatory reasons
(Grier 1975).
Sunning (basking) behavior is widely studied among ectothermic
vertebrates (Heath 1970, Greenberg 1980), but the effects of incident
radiation and its importance in energetics are often overlooked in
studies of endotherms. In environments that impose energetic stress,
the tapping of auxiliary energy sources such as insolation could re
duce an endotherm's costs of thermoregulation. Sunning behavior has
been reported within several diverse avian orders including Passeri
formes (Hauser 1957, Lanyon 1958), Cuculiformes (Ohmart and Lasiewski
1971, Grier 1975), Ciconiiformes (Clark 1969, Curry-Lindahl 1970,
Kahl 1971), Psittaciformes (Cade 1973), Falconiformes (Clark 1969,
Hatch 1970, Cade 1973, Grier 1975), and Pelecaniformes (Snow 1966,
Clark 1969, Curry-Lindahl 1970).
The fact that birds can take advantage of solar radiation to
reduce the metabolic costs of endothermy has been demonstrated for
several species of diverse origins (Hamilton and Heppner 1967a,
1967b, Morton 1967, Lustick 1969, 1971, Heppner 1969, 1970, Lustick


3
et al. 1970, De Jong 1976). Ohmart and Lasiewski (1971) have shown
that Roadrunners (Geococcyx californianus) utilize solar radiation in
conjunction with nightly hypothermia to reduce metabolism, and there
fore food intake. A similar mechanism has been proposed for the
Turkey Vulture (Cathartes aura) by Heath (1962). Nocturnal tempera
ture decline may be widespread among avian species (Irving 1955).
The role, if any, that sunning plays in most of these cases is largely
unknown.
The capacity to reduce energetic requirements through insolation
absorption is enhanced by dark pigmentation (Hamilton and Heppner 1967a&b,
Heppner 1970, Lustick 1971). An increase in surface area exposed to
the sun (such as with a spread-winged posture) increases overall
solar energy interception, i.e., sunning. At ambient temperatures
below thermoneutrality insolation may, therefore, be an aid to thermo
regulation for many species, allowing them to reduce rates of heat
loss (thermal conductance). However, at higher ambient temperatures
insolation may tax the abilities of darkly pigmented birds to dissipate
heat produced by metabolism and to maintain normal body temperatures
in warm, humid environments such as Florida (Wunder 1979).
Cormorants and anhingas are excellent subjects for a study exam
ining the relationships among insolation, spread-winged behavior, and
thermoregulation because of their conspicuous use of the spread-winged
posture and their predominately black coloration. In addition to
overall dark coloring, the Anhinga has a subalular apterium of darkly
pigmented skin (George and Casler 1972), which potentially has a
thermoregulatory function similar to the interscapular apterium of
the Roadrunner (Ohmart and Lasiewski 1971). Spread-winged behavior


4
within these two families has been considered as wing drying, as both
anhingas and cormorants have water absorbing plumage (Rijke 1968,
easier 1973) that allows water to penetrate air spaces in the plumage,
reducing buoyancy and facilitating underwater prey capture.
This study examines the thermoregulation of the Anhinga and the
Double-crested Cormorant in the laboratory, and their behavior in the
field to determine 1) the effects of insolation on energetics and
behavior, and 2) whether their energetics and behavior support either
a wing drying or thermoregulatory function for spread-winged behavior.
Also included is a look at spread-winged behavior and thermoregula
tion in the Flightless Cormorant (previously Nannopterum harrisi, now
Phalacrocorax harrisi, Mayr and Cottrell 1979), which shares this
behavior with its cormorant relatives.


CHAPTER TVJO
METHODS AND STUDY SITES
Field Studies
Anhingas and Double-crested Cormorants
I observed Anhinga behavior at Lake Alice, 29 39' N lat.,
82 20' W long, (fresh water) in Gainesville, Florida, and Double-
crested Cormorants at Seahorse Key, 29 34' N lat., 83 4' W long.,
Florida (salt water), Lake Wauberg, 29 38' N lat., 82 34' W long.,
near Micanopy, Florida (fresh water), Bivens Arm, 29 38' N lat.,
82 18' W long., in Gainesville, Florida (fresh water), and Lake
Alice. Observations were made at all times of the day and in all
seasons during 1978-1979 (Anhingas) and 1979-1980 (cormorants). At
5-min intervals I recorded (1) the amount of time that each bird
under observation spent performing activities such as wing spreading,
preening, foraging, and guiar flutter, (2) the orientation of each
individual with respect to wind and sun, (3) ambient (shade) tempera
ture, and (4) the intensity of solar radiation measured with an
Eppley precision spectral pyranometer. This instrument measures both
direct and diffuse radiation in a 180 field and was aimed directly at
the sun to assess the radiation levels available to sunning birds.
Unless otherwise indicated, all discussions of solar radiation in this
study refer to solar radiation measured in this fashion. The behavior
recorded at the end of each interval was considered one observation
5


6
of that behavior. If the individual under observation exhibited
more than one behavior during the 5-min interval, I recorded the
fraction of the interval devoted to each behavioral category as
no. of total observations of the behavior ^ -jqq
total no. of observations
under each specific set of environmental conditions. For example, if
I made 20 observations of Anhingas while the intensity of solar
O
radiation was below 222 W/m and ambient temperature (T ) was between
0 and 5C, and in 2 observations birds were preening, then preening
was considered to represent 10% of the behavior observed under those
environmental conditions. I then examined these data for correlations
between the frequency of certain behaviors and environmental conditions.
Because Double-crested Cormorants are nonterritorial (unlike
Anhingas) and moved more frequently and over longer distances than did
Anhingas, it was frequently impossible to follow individuals for ex
tended time periods. This inability to follow individuals made it
difficult to take accurate time budget data. Therefore, while observ
ing cormorants, in addition to the data on individual birds (described
above), I also recorded the following data on the entire visible popu
lation: number of individuals in the water, number perched out of
the water, number engaged in spread-winged behavior, and number of
individuals guiar fluttering. These data recorded on the population
as a whole were used to determine the frequencies of the behaviors
listed above.


7
Flightless Cormorants
I observed Flightless Cormorant behavior in the vicinity of
Caleta Tagus, 0 15' S lat., 91 22' W long., Isla Isabela, Galapagos,
Ecuador, during July and August 1980. Observations were made at all
times of the day and night during the study period. At 5 minute
intervals I recorded (1) behavior, (2) orientation to sun and wind,
(3) shade T and (4) solar intensity as measured with a black-body
a
thermometer constructed from a 5 cm by 5 cm piece of wood with a groove
into which a thermometer was fitted and covered with a fine sheet of
brass, spray-painted black. Occasional measurements of direct solar
radiant intensity were made with an Eppley precision spectral
pyranometer.
In addition, body temperatures were measured simultaneously with
behavioral and environmental data using miniature, temperature-
sensitive, radio transmitters (Model T, Mini-mitter Co., Indianapolis,
Indiana) that emit a train of pulses, the frequency of which is tem
perature dependent. These were received with a Lafayette walkie-
talkie. Transmitters were calibrated using a temperature controlled
water bath with a thermostat and a mercury and glass thermometer.
Cloacal T^'s were also taken with a Schultheis thermometer whenever
it was possible to do so without interfering with behavioral observa
tions. Using these techniques, I monitored Tb's during the per
formance of various behaviors such as spread-winged behavior, foraging,
and guiar flutter, as well as for 24 hour cycles to determine normal
daily rhythms.


8
Laboratory Studies
1 hand-raised four Anhingas (2 adult females with an average
weight of 1.12 kg; 2 juvenile females averaging 0.93 kg), housed them
in an outdoor aviary in which they were exposed and acclimated to
Gainesville weather conditions, and maintained them on a diet of
whole smelt and minnows. I borrowed five Double-crested Cormorants
(2 adults and 3 juveniles, mean weight = 1.33 kg) from the Suncoast
Seabird Sanctuary in Indian Shores, Florida, and maintained them as
with the Anhingas. I measured temperature-specific rates of metabolism
for each individual over a range of ambient temperatures (-10 to
45C) in an open flow system employing a Beckman G2 or Applied Electro
chemistry S-3A oxygen analyzer. Measurements were made in a constant-
temperature chamber with a volume of 326 1, using flow rates of 4.4 to
6.8 1/min. Flow rates were measured with a Brooks R215-B rotameter
after the removal of CO,, and water; all values were corrected to
STPD. Chamber temperatures were measured with a mercury and glass
thermometer guarded with a radiation shield suspended inside the
chamber. I measured metabolic rates when the subjects were in a
postabsorptive state under the following environmental conditions:
(1) at night without incident radiation, (2) during the day without
incident radiation, (3) daytime with incident radiation = 450 watts/
2 p o
meter (W/m ), (4) daytime with incident radiation = 700 W/m .
Simulated solar radiation was provided by a pair of white heat lamps
(either 2 Westinghouse 125 W lamps or 1 125 W lamp and 1 GE 250R40/1
250 W lamp) positioned to shine through a 25 cm x 25 cm piece of Pyrex
(which transmits near ultraviolet) fitted into the chamber lid. The


9
subjects perched roughly 45 cm behind the Pyrex window, and radiation
intensities were measured at this distance using an Eppley precision
spectral pyranometer. I measured resting metabolism only after
chamber temperature had stabilized, equilibrium conditions had been
satisfied (Christensen 1947, Heusner 1955), and VO2 had stabilized.
I monitored the bird's deep body temperatures using Mini-mitters,
which were coated with beeswax and fed to the subjects. I also
measured cloacal temperatures after each metabolic trial using a
Schultheis thermometer. Both methods of measuring body temperature
yielded statistically similar results and are used interchangeably
throughout the text.
Mean basal rates of metabolism were estimated from independent
measurements of the minimal rate of oxygen consumption within the zone
of thermoneutrality. I estimated minimal thermal conductances (C )
m
from the relationship
Cm = iV 0)
using simultaneous measurements of metabolic rate (VC^), body tempera
ture (T^), and ambient temperature (T ) at temperatures below thermo
neutrality. I compared basal metabolic rates and minimal conductances
measured under control conditions (without incident radiation) with
those measured under experimental conditions (with incident radiation)
and with those expected from body mass. "Expected" values for basal
metabolic rate were calculated from the predictive equations derived
from Aschoff and Pohl (1970):


10
V02/M = 4.0 M"0,27 (2)
for a nonpasserine in the quiescent phase of the diurnal cycle, and
V02/M = 5.14 M"0'27 (3)
for a nonpasserine in the active phase of the diurnal cycle, where
. 3
V02/M is basal metabolic rate in cm O^/g h and M is mass in grams.
Expected values for mean minimal conductances were calculated from the
predictive equations of Aschoff (1980):
C/M = 0.947 M"0-583 (4)
for a nonpasserine in the quiescent phase of the diurnal cycle, and
C/M = 0.946 M~484 (5)
for a nonpasserine in the active phase of the diurnal cycle, where
3
C/M = "wet" thermal conductance in cm 02/g h C (Mcnab 1974), and
M is in grams.
To determine the effect of water on thermoregulation (during
daytime) each individual was placed unrestrained in water 16 cm deep
(to midbreast in Anhingas, to lower breast in cormorants) inside the
environmental chamber, at ambient temperatures of from 12 to 35C for
two hour time periods, while metabolic rate and body temperature were
monitored as described above. Equilibria! metabolic rates and T^'s
recorded under these conditions were compared with those recorded in


11
dry individuals. Before placing them in the chamber with water, each
bird was submerged fully (except for the head) for at least one
minute, until the plumage was saturated.
I also estimated evaporative water loss (EWL) to assess the in
fluence of insolation both within and above the thermoneutral zone.
Evaporative water loss was measured during metabolic trials using
techniques similar to those described in Lasiewski et al. (1966).
Individual birds stood in the environmental chamber on a trap filled
with paraffin oil and covered with hardware cloth to capture any
excrement eliminated during the trial, thus removing a possible source
of error in the calculation of EWL.
The stream of air passing through the metabolic chamber was
analyzed for water content immediately after leaving the chamber.
I measured total water content gravimetrically by passing the air
stream through a drying tube filled with dessicant (Drierite) for a
known time interval and at a constant flow rate. I then calculated
the evaporative water loss from the subject by subtracting out the
water content of room air, which was measured gravimetrically by
drying room air in a fashion similar to that used on the sample coming
from the environmental chamber. At high environmental temperatures
inside the chamber the stream of air coming from the chamber was
heated with a heating coil (controlled with a rheostat) wrapped
around the tubing housing the air stream to prevent condensation be
fore the drying tube. As with metabolic measurements, measurements
of EWL were made only after equilibrium conditions had been satisifed
(Lasiewski et al. 1966). After calculating EWL over a range of
chamber temperatures for each species and under each set of


12
environmental conditions (daytime without incident radiation, daytime
2
with incident radiation = 450 W/m daytime with incident radiation =
o
700 W/m ), I compared the rates of evaporative water loss under each
set of environmental conditions both within and between species.


CHAPTER THREE
RESULTS: FIELD OBSERVATIONS
Anhingas
My field observations on Anhingas are summarized in Table 1.
Behavior in general, and wing spreading in particular, strongly depends
on weather. Anhingas exhibit spread-winged postures much more fre
quently during cool than during warm weather. The frequency of this
behavior is negatively correlated with air temperature (Fig. 1, r =
-0.85, p < 0.01, 9df) but positively influenced by the intensity of
2
solar radiation (Table 1). When the intensity was less than 222 W/m
(cloudy days with mostly diffuse radiation), spread-winged behavior
constituted only 3.8% of the observed behavior, but it accounted for
2
46.5% of the observed behavior when solar intensity exceeded 222 W/m
(T < 30C in both cases, p < 0.001, test for the equality of two
percentages, Sokal and Rohlf 1969). When solar radiation levels were
low (e.g., during early evening), birds just emerging from the water
often fanned their wings and tails continuously (fan drying, Table 1)
rather than holding them steady (as was usually the case).
Thirty-four percent of the observed spread-winged behavior in
volved birds that appeared to be wet, 49% involved individuals that
appeared to be dry, and 17% involved individuals whose plumage con
dition could not be determined. Wet birds usually held their wings
completely outstretched, whereas dry individuals often held them
closer to the body or occasionally drooped at their sides. During
13


Table 1. Time budgets for Anhingas under various environmental conditions
Environmental
Conditions
Behavioral
Frequency (%)
Solar
Intensity
Ta
Sunning
Sun &
Preen
Sun
Total
Preen
Perching
Guiar
Flutter
Swim
Heat
Conserving
Posture
Fan
Dry
Total
Observ.
<222 W/m2
<30C
1.6
2.2
3.8
13.0
43.6
0.0
5.4*
31.8*
2.5*
387
>222 W/m2
<30C
40.2*
6.3
46.5*
9.7
22.1
2.6
16.9
1.6
0.0
860
<222 W/m2
>30C
10.0
0.0
10.0
28.0
32.5
0.0
16.9
0.0
0.0
25
>222 W/m2
>30C
8.8
0.0
8.8
17.1
32.0
23.8*
18.2
0.0
0.0
111
Total Time Budget
26.3
4.6
30.9
11.4
29.1
3.5
14.0
9.4
1.1
1383
Behavioral Frequency = percent of total observations recorded under those conditions in which that specific
behavior was observed (see text).
Sunning refers to spread-winged behavior without accompanying preening.
Sun & Preen refers to spread-winged behavior accompanied by preening.
*Statistically significant difference (Test for the equality of two percentages, Sokal and Rohlf 1969).


Figure 1. Relationship between frequency of spread-winged behavior and ambient temperature
for Anhingas and Double-crested Cormorants


BEHAVIORAL FREQUENCY (%)


17
spread-winged behavior Anhingas commonly held their wings and body
perpendicular to the direct beam incident solar radiation, flattening
out when the sun was high, standing erect when the sun was low.
Concordantly, most birds displaying spread-winged postures oriented
with their backs to the sun; in only 16 or 3.4% of 468 observations
did they orient differently. In these 16 cases, all had wet plumage,
most (14/16) were facing into the wind, and most (14/16) were observed
when the intensity of solar radiation was less than 222 W/m The
frequency with which Anhingas oriented with back to the sun was also
negatively correlated with Ta (r = -0.83, p < 0.01, 9df).
Anhingas oriented quite differently while heat stressed (exhibit
ing guiar flutter), facing into the sun 53% of the time, obliquely
(side to the sun) 36% of the time, and away from the sun only 11% of
the time (55 observations). This orientation is statistically non-
random (x^ = 43.12, p < 0.001, 2 df). Guiar flutter occurred when Ta
was between 25 and 30C, but only when the intensity of solar radia-
2
tion exceeded 700 W/m All other observations of guiar flutter oc
curred when Ta was above 30C. Guiar flutter was significantly more
common at high levels of solar radiation (Table 1) and was never
accompanied by a spread-winged posture. Orientation during guiar
flutter (predominately facing the sun) is similar to that during ex
posure to heavy winds in which individuals usually face into the wind,
thereby exposing a minimum of surface.
Anhingas spent 14.0% of the time foraging in the water during
the study. The bulk of their time (comprising over 80% of the total
observations) was spent perching (preening, sunning, or just sitting).
They spent significantly less time in the water when T, was below 25C
a


18
(10.8% of 783 total observations) than when it exceeded 25C (23.8%
of 600 total observations, test for the equality of two percentages,
p < 0.001).
After leaving the water Anhingas shook excess water from their
plumage, and almost invariably spread their wings. However, they also
displayed spread-winged postures with dry plumage before entering the
water. Seventy-one percent of all trips to the water, and 90% of all
2
such trips when solar radiation exceeded 222 W/m were preceded by
spread-winged behavior. This value increases to 94% if one considers
only those occasions when Ta was below 30 and solar radiation exceeded
a
222 W/m2.
At low Ta, both in the field and during metabolic trials, Anhingas
commonly folded their necks in a tight S-shape and pressed them tightly
against the upper body. Occasionally, individuals assumed a posture
in which the head and neck were laid along the back and covered with
the wings. Both postures should reduce thermal conductance by reducing
surface area and I, therefore, labelled them hzat con^nAvlnq po&WiQA
(Table 1).
Double-Crested Cormorants
Field observations of individual cormorants are summarized in
Table 2; data taken on the population as a whole are presented in
Table 3. Unlike Anhingas, the behavior of Double-crested Cormorants is
not especially sensitive to weather. The frequency of spread-winged
behavior in these cormorants is negatively correlated with ambient
temperature (Fig. 1) (r = -0.54, p > 0.05, 5 df) but not as dramatically
as in Anhingas. As with Anhingas, spread-winged frequency is positively


Table 2. Frequencies for the behavior of individual Double-crested Cormorants under various
environmental conditions
Environmental
Conditions
Behavioral Frequency (%)
Solar
Radiation
W/rrr
(C)
Total
Observations
Spread-
winged
Behavior
Spread
winged
Preening
Preening
Perching
Guiar
Flutter
Head
Under
Wi ng
Delta
Posture
>222
<20
337
4.3
0.3
30.7
49.5
12.2
3.0
0.0
<222
<20
122
0.8
0.0
23.4
70.1
0.0*
5.7
0.0
>222
>20
276
1.6
0.0
27.4
64.8
4.7
0.0*
1.5*
<222
>20
39
0.0
0.0
33.8
63.6
0.0*
2.6
0.0
774
2.6
0.1
28.5
58.8
7.1
2.3
0.5
*Significantly different (P < 0.05) from the other percentages in the column (test for the equality of
two percentages (Sokal and Rohlf 1969)


Table 3. Frequencies for the behavior of a population of Double-crested Cormorants under various
environmental conditions
Environmental Conditions
Solar Intensity
(W/rri2)
Ta
(6)
Total
Observations
Spread-winged3
Behavior
Delta3
Posture
Guiar3
Flutter
Percent in
Water
Percent out
Water
<222
<20
825
2.0
0.0
0.0
7.8
92.2
>222
<20
2,932
2.7
0.0
6.6
34.0
66.0
<222
>20
586
0.0
0.0
0.0
78.5b
21 5b
>222
>20
1,179
1.2
4.5
17.1
22.2
77.8
aTaken as a percent of the behavior recorded on land (not including observations of birds in the water)
bBiven's Arm (see text)


21
influenced by solar intensity. Linear correlation between spread
winged frequency and solar intensity reveals the relationship:
y = 1.88 + O.OOlx (r = 0.36, p > 0.05, 5 df). However, spread-winged
behavior never exceeded 4.1% of the observed behavior under any en
vironmental conditions and accounted for only 2.3% of the total ob
served behavior of cormorants on land (Table 3).
Spread-winged behavior was only observed in birds with (apparently)
wet plumage and was often accompanied by wing fanning. Unlike Anhingas,
cormorants did not exhibit spread-winged behavior before entering the
water and did not display any but full spread-winged postures, with
wings fully extended.
Orientation with respect to the sun during spread-winged behavior
was nonrandom (y^ =11.11, p < 0.005, 2 df) with 39.2% of the birds
facing away from the sun, 43% facing obliquely to the sun, and 17.8%
facing into the sun. Unlike Anhingas, Double-crested Cormorants did
not hold their wings and bodies perpendicular to the sun during spread
winged behavior but always stood erect, never flattening out when the
sun was high. Orientation with respect to the wind was also nonrandom
(X^ = 51.88, p < 0.005, 2 df) with 17.9% facing away from the wind,
26.1% facing obliquely, and 56% facing into the wind during spread
winged behavior.
As with Anhingas, cormorants also oriented differently during
guiar flutter than when engaged in spread-winged behavior. Ninety per
cent of all individuals engaged in guiar flutter were facing the sun.
Guiar flutter was observed at ambient temperatures as low as 12C with
incident radiation in excess of 1,000 W/m and was only observed once


22
when radiant intensity was below 222 W/m^ (T > 20C). Guiar flutter
a
was never accompanied by a spread-winged posture.
Two behaviors were observed in Double-crested Cormorants that
were not observed in Anhingas. One such behavior was displaying spread
winged postures while floating on the water's surface, which was ob
served on 4 occasions in cormorants at Lake Wauberg. This variation
of spread-winged behavior has also been reported for European Cormorants
(J?. carbo) and for Shags (£_. aristotel is) (Fry 1957). The second
behavior unique to cormorants was, again, a variation of spread-winged
behavior, in which the wings were not held straight out to the side,
but were bent at the elbow and folded down into a delta-winged posture
commonly observed in storks (Kahl 1971). This behavior was only ob
served at Seahorse Key (where much of my hot weather data were recorded)
O
with Tfl in excess of 30C and solar radiant intensity above 900 W/m
and was usually accompanied by guiar flutter. Orientation during this
behavior was invariably facing into the sun.
Cormorants spent 32.3% of their time in the water during the
study, but, as with Anhingas, the bulk of their time was spent perch
ing. Although cormorants spent more time in the water when T. exceeded
a
20C (28.3% of the observations recorded with Ta < 20C as compared to
40.9% of the total observations with Tfl > 20C, p < 0.001, test for the
equality of two percentages), the significance of this difference may
be entirely due to sampling error, as many of the observations of
birds at ambient temperatures above 20C were recorded at Biven's Arm,
where large rafts of cormorants were commonplace (apparently) irre
spective of weather. This rafting behavior was not observed at the
other study sites and may have unfairly skewed the data with respect


23
to time spent in the water. However, this rafting behavior, and my
subjective observations, suggest that cormorants spend longer time
periods in the water at a stretch, and as a percentage of the day,
than do Anhingas.
Flightless Cormorants
A time budget summarizing the behavioral data for Flightless
Cormorants is presented in Table 4. I never observed Flightless
Cormorants exhibiting spread-winged postures while having dry plumage;
true spread-winged behavior was exhibited only after the animal had
left the water. Only once was a cormorant observed to leave the water
without subsequently displaying this behavior. As with Anhingas and
Double-crested Cormorants, I never observed spread-winged behavior in

Flightless Cormorants accompanied by guiar flutter. A drooped-winged
posture was observed in dry individuals, often while tending the nest.
This posture differed from true spread-winged behavior in that the
wings were not fully extended out to the sides but were held loosely
downward at the side of the body. Drooped-winged postures were accom
panied by guiar flutter 811 of the time and were most commonly ob
served when the sky was cloudy and ambient temperatures were moderate
(Tables 5, 6). Guiar flutter was most commonly observed (as expected)
during sunny conditions and at high ambient temperatures (Table 5, 6).
Spread-winged behavior was most commonly observed when the sky
was cloudy and ambient temperatures were high (Tables 5, 6). Linear
correlation between the frequency of spread-winged behavior and
ambient temperature reveals a positive relationship (just the opposite
of Anhingas) with r = 0.92 (p < 0.05). Correlation between the


Table 4. Time budgets for Flightless Cormorants on and off of the nest
Behavioral Frequency (%)a
Spread
winged
S-W &b
Preen
Preen
Sitting
Standing
Guiar
Flutter
Drooped-
winged
Fan
Dry
Head
Under
Wi ng
0therc
Total
Observa
tions
Tending
Nest
1.3
0.6
4.0
36.8
15.8
37.5
6.7
2.2
5.5
3.4
806
Off of
Nest
5.8
11.2
38.7
2.0
29.1
9.4
0.8
0.0
7.0
2.3
395
Combined
2.8
4.1
15.4
25.3
20.2
28.4
4.7
0.2
5.9
3.0
1201
aBehavioral Frequency = percent of total observations devoted to that particular class of behaviors.
Behavioral categories are not mutually exclusive.
^Preening while spread-winged
c0ther includes interacting socially, arranging nest material, shaking off, walking, etc.


Table 5. Some behavioral frequencies for Flightless Cormorants exposed to different amounts of
sky cover
Behavioral Frequency (%)a
Sky Cover
Spread-winged
Behavior
Drooped-winged
Posture
Guiar
FI utter
Total ,
Observations0
Clear
8.94
3.63
52.52
475
Cloudy
12.98
15.58
44.15
39
Shadec
2.95
4.43
8.13
271
aBehavioral Frequency = percent of total observations recorded under those sky conditions
devoted to that specific behavior
bTotal Observations refers to all observations recorded for birds on land under those specific
sky conditions
c
Shade refers to observations recorded when the subject was in the shade or when the sun
had set.


Table 6. Some behavioral frequencies for Flightless Cormorants exposed to different ambient
temperatures
Behavioral Frequency (%)a
Ta
Spread-winged
Behavior
Drooped-winged
Posture
Guiar
Flutter
Total
Observations
20-22.5C
8.25
0.61
10.39
164
22.5-25C
8.09
11.74
38.25
247
25-27.5C
11.63
10.34
45.68
232
27.5-30C
12.32
0.91
65.75
no
aBehavioral Frequency = percent of total observations recorded under those ambient temperatures
devoted to that specific behavior
^Recorded for cormorants on land in that specific temperature range


27
duration of spread-winged sunning episodes (the length of time the
wings were extended during a bout of sunning) of individual cormorants
and ambient temperature reveals a negative relationship with r = -0.28
(p > 0.05). Linear correlations between the duration of "sunning"
episodes and black-body temperature reveals a negative relationship
with r = -0.35 (p < 0.05).
Orientation of individual birds with respect to the sun during
spread-winged behavior was nonrandom (p < 0.01), with the primary skew
in the data favoring orientation with back to the sun; 46% of all ob
servations of "sunning" birds were of individuals orienting with back
to the sun. Unlike Anhingas and Double-crested Cormorants, however,
orientation of Flightless Cormorants during guiar flutter was virtu
ally identical to orientation during spread-winged behavior, with
the primary skew in the data favoring orienting with back to the sun
(42%). In fact, orientation of cormorants without regard for behavior
was nonrandom, again with the primary skew in the data favoring orien
tation with back to the sun (42%). Therefore, orientation in Flight
less Cormorants does not depend (apparently) on behavior; these cormor
ants preferentially orient with back to the sun and avoid orientations
in which they face into the sun. Orientation of all birds observed,
however, did vary with sky cover (Table 7).
Orientation to the wind during spread-winged behavior was non-
random (p < 0.05), with the primary skew favoring facing into the wind
(42%). Only 13.5% of the birds observed "sunning" were facing with
back to the wind. Orientation of Flightless Cormorants to the wind,
regardless of behavior, was again nonrandom (p < 0.05), and again the
primary direction of data skew was in favor of facing into the wind,


Table 7. Orientation of Flightless Cormorants under different sky conditions.
Orientation
(% of total observations)
Sky Cover
Back to the Sun
Side to the Sun
Front to the Sun
Total Observations
Cl ear
48
38
14
492*
Partly Cloudy
62
36
2
46*
Overcast
26
55
19
230
Sun Set
42
56
2
57*
*Nonrandom Orientation (Chi-square, 2 df)


29
regardless of wind strength (34.7% of observations with wind calm to
light, 46.7% of observations with wind moderate to heavy were of birds
facing into the wind).
The amount of time spent foraging v/as impossible to estimate ac
curately for these cormorants, as they frequently swam out of sight and
often left the water and "sunned," or simply rested far from the nest
(and from view). However, foraging represented at least 9.8% of the
total behavior observed (based on observations of subjects swimming in
view). Eleven foraging bouts witnessed start to finish lasted from 5
to 43 minutes in length (average = 18.4 minutes).
Body temperatures measured for Flightless Cormorants during various
behaviors and under different conditions are presented in Table 8. The
T^'s averaged higher during the day than at night, while guiar flut
tering as opposed to while not guiar fluttering (day only), before
entering the water as opposed to leaving the water, and before exhibit
ing spread-winged behavior as opposed to after "sunning" (Table 8).
The lowest body temperatures recorded were taken at approximately 0330
(mean Tfa = 39.2C 0.32 SD).


Table 8.
Body temperatures
a
for flightless conditions under various conditions
Day
Night
Guiar
FI utter
Not Guiar
Fluttering
Entering
Water
Leaving
Water
After
Spread-winged
Behavior
Range of
38.7-41.3
38.7-40.2
39.8-41.3
38.7-41.2
40.3-41.2
39.0-41.2
39.4-40.1
Mean
40.11
39.47
40.36
39.89
40.70
39.90
39.72
SD
0.52
0.47
0.29
0.58
0.47
0.64
0.34
Number of
Measurements
54
18
25
29
4
13
9
ain C


CHAPTER FOUR
LABORATORY RESULTS
Energetics: Day versus Night
The relationships among metabolic rate, body temperature (T^), and
ambient temperature for the metabolic trials featuring birds with dry
plumage and no incident radiation are presented in Figures 2 (Anhingas,
measured during the day), 3 (Anhingas, measured during the night),
4 (cormorants, measured during the day), and 5 (cormorants, measured
during the night).
Anhingas have lower than expected basal metabolic rates and higher
than expected thermal conductances; Double-crested Cormorants have
higher than expected thermal conductances and metabolic rates regard
less of circadian phase (Table 9). Metabolic rates measured below
thermoneutrality during the day averaged 30-32% higher (Anhingas) and
15-30% higher (cormorants) than those measured at night. Body tempera
tures averaged higher during the day than at night (Table 9) and gen
erally rose with ambient temperature suggesting a tendency for heat
storage (Figures 2-5).
The slopes of the regression lines fitting the data presented in
Figures 2-9 cannot be used as estimates of thermal conductance because
these lines do not extrapolate to body temperature, indicating that
Anhingas and Double-crested Cormorants mix physical (i.e., postural
and plumage adjustments) with chemical thermoregulation (i.e.,
metabolism) at temperatures below thermoneutrality (McNab 1980).
31


Figure 2. Relationships among metabolic rate, body temperature, and ambient temperature
for 4 Anhingas as measured during the day


RATE OF METABOLISM (cm3 02/g hr)
30DY


Figure 3
i
j
J
i
I
j
j
1
f
i
1
I
I
Relationships among metabolic rate, body temperature, and ambient temperature
for 4 Anhingas as measured during the night J
I


RATE OF METABOLISM (cm3 02/ghr)
o


Figure 4. Relationships among metabolic rate, body temperature, and ambient temperature
measured for 5 Double-crested Cormorants during the day


RATE OF METABOLISM (cm3 02 /g-hr)
i- 4
O
CL
UJ
I-
>"
O
o
m
i
CO
--0


I
¡
i
1
1
I
i
i
I
i
I
i
I
¡
Figure 5. Relationships among metabolic rate, body temperature, and ambient temperature as
measured for 5 Double-crested Cormorants during the night


RATE OF METABOLISM (cm3 Og/ghr)
30DY TEMP (C)


Table 9. Thermoregulation in Double-crested Cormorants and Anhingas: Day versus night
Species
BMR
(% Expected)
Tb(in)
^b(below)
Ti
C
m
(% Expected)
Phalacrocorax auritus
Day
0.85
115
41.2
40.7
21 .0
0.0368
126
Night
0.74
128
40.2
39.7
17.5
0.0320
224
Anhinqa anhinqa
Day
0.60
75
39.9
39.7
29.5
0.0454
138
Night
0.55
89
39.1
38.0
26.0
0.0364
221
o
BMR is basal metabolic rate in cni 02/g-h
l Expected is a comparison of measured metabolic rates with expected values (equations 2 and 3)
Tb(in) is body temperature (C) measured in thermoneutrality
^b(below) boc*y temperature (C) measured below thermoneutral ity
T^ is the lower limit of thermoneutrality (C)
C|n is minimal thermal conductance in cm 02/g-h-C
% Expected Cm is a comparison of minimal thermal conductances with expected values (equations 4 and 5)


41
Energetics: Incident Radiation
The relationships among metabolic rate, body temperature, and
ambient temperature during exposure to incident radiation are presented
in Figures 6-9. Mean body temperatures and basal rates of metabolism
in both species were higher under incident radiation, possibly reflect
ing a higher activity level (Table 10). Thermal conductances were
lower under incident radiation, especially in Anhingas (Table 10).
Cormorants never displayed spread-winged postures with or without
incident radiation unless first fully soaked with water, and then only
for short intervals (rarely more than 10 min). No reduction in meta
bolic rate was observed during these sunning episodes. Although
cormorant thermal conductances were slightly lower than control values
2
when the birds were exposed to 700 W/m incident radiation, and
o
slightly higher when exposed to 450 VJ/m (Table 10), neither differ
ence was statistically significant. A comparison of the regression
lines fitting the three data sets (Fig. 10) indicates a slight but non
significant (Table 10) reduction in slope under incident radiation.
Consequently, rates of metabolism measured below T-j with incident
radiation are virtually 100% of those measured without incident radia-
2 2
tion during exposure to 450 W/m and 90% under 700 W/m incident
radiation. Under incident light the extrapolated T^ (x-intercept)
is well above the actual T^ (Table 10), indicating that the added
physical factor (incident light) is being absorbed and is increasing
the physical component of thermoregulation. However, there is no
statistical difference in either thermal conductance or regression
slopes between the experimental and control data for Double-crested
Cormorants. Additionally, the lower limit of thermoneutrality (T^)


Table 10. A comparison of the energetic responses of Double-crested Cormorants and Anhingas to
variation in ambient temperature and incident radiation
Species
Treatment
BMR
Tb
cm
cf
r
Ti
Extrap.
Tb
Phalacrocorax auritus
no lamp
0.85
40.70
0.0368
0.0281
-0.90
21.0
50.92
450 W/m2
0.91
41 .11
0.0373
0.0230
-0.50
23.0
62.17
700 W/m2
0.95
41.36
0.0352
0.0176
-0.81
19.5
73.18
Anhinqa anhinqa
no lamp
0.60
39.70
0.0454
0.0340
-0.92
29.5
47.06
450 W/m2
0.70
39.97
0.0290
0.0160*
-0.79
17.5
61 .84
700 W/m2
0.72
40.05
0.0272
0.0106*
-0.59
17.0
84.98
o
BMR is basal metabolic rate in cm O^/g-h
Tb is body temperature measured at temperatures below thermoneutrality in C
3
is minimal thermal conductance in cm 02/g-h-C
3
Cf is a measure of thermal conductance determined by the method of least squares (cm 02/g-h-C)
is the lower limit of thermoneutrality (C)
Extrapolated Tb is the Tb to which the regression line fitting the data extrapolates
O
Experimental class differs significantly from control (Mann-Whitney U)

Experimental class differs significantly from control


Figure 6. Relationships among metabolic rate, body temperature, and ambient temperature
as measured for 4 Anhingas under 450 W/rrr incident radiation


RATE OF METABOLISM (cm3 02/ghr)
-pi
-p=>
BODY TEMP (C)


Figure 7. Relationships among metabolic rate, body temperature, and ambient temperature
as measured for 4 Anhingas under 700 W/nr incident radiation


RATE OF METABOLISM (cm3 02/ghr)


Figure 8. Relationships among metabolic rate, body temperature, and ambient temperature as
measured for 5 Double-crested Cormorants under 450 W/m^ incident radiation




Figure 9.
Relationships among metabolic rate, body temperature, and ambient temperature as
measured for 5 Double-crested Cormorants under 700 W/m^ incident radiation


RATE OF METABOLISM (cm302/ghr)
o
o
cn
o


Figure 10
Comparison of the regression lines fitting control and experimental data sets for
Anhingas and Double-crested Cormorants
I
i
l
1
i
l


METABOLIC RATE (cm3 02/g-h)
ENVIRONMENTAL TEMPERATURE (C)


53
under incident radiation is virtually unchanged from that measured
under control conditions for this.._species (Table 10).
These results contrast with those for Anhingas. In the aviary I
observed spread-winged behavior in both wet and dry anhingas almost
daily, especially when ambient temperature was below T^. Attempts to
elicit this behavior in the experimental chamber were frustrated until
2
the incident radiation was increased to exceed 250 W/m indicating that
Anhingas have a threshold below which spread-winged sunning may not be
profitable. With incident radiation above this threshold and ambient
temperature below T^, experimental subjects generally extended their
wings with the onset of illumination, and often kept the wings spread
for an hour or more. Occasionally spread-winged postures were observed
above T^, but only for'brief intervals. ~ Anhingas typically held the
wings perpendicular to the incident light, presumably to take full ad
vantage of the heat-absorbing qualities of the dark plumage (Lustick
1980) and erected back and wing feathers to expose areas of the back.
With the onset of sunning, VC^ steadily declined until a new equi
librium was reached and spread-winged behavior ceased. This pattern
of continuously declining VO2 was never observed in trials in which
spread-winged behavior was absent. One individual frequently v/ould not
sun or would not hold the spread-winged posture for longer than a
minute or two; this bird maintained metabolic rates and thermal con
ductances equivalent to, or higher than, those observed under control
conditions. This suggests that the spread-winged posture is necessary
to reduce VO^ significantly in the presence of incident radiation. In
birds displaying spread-winged postures, exposure to incident light
significantly reduced both thermal conductances and the slopes of the


54
regression lines (Table 10). The rates of metabolism measured under
experimental conditions at temperatures below T-j averaged 62% of those
2
measured under control conditions during exposure to 450 W/m and
only 56% of control under 700 W/m^. The reduction in thermal con
ductance during exposure to insolation enables the Anhinga to lower T-|
by 12 to 12.5C (Table 10).
Evaporative Water Loss
The relationships between log evaporative water loss and ambient
temperature for birds with dry plumage under varying intensities of
incident radiation are presented in Figure 11 (Anhingas) and Figure 12
(cormorants). Semi log plots were adopted so that these curvilinear
relationships could be transformed into linear relationships for the
purpose of comparing them statistically. In each species there is a
significant difference in the rate of evaporative water loss (p < 0.05,
Analysis of Covariance) between the measurements made without incident
radiation and those with radiation. However, there are no significant
differences (p > 0.05) between the responses to the two different in
tensities of incident radiation. There is little difference in the
rate of evaporative water loss between the cormorant and Anhinga when
radiation is present, but in the absence of incident radiation the
Anhinga has a lower rate of water loss (Figs. 11, 12). Because the
Anhinga has a lower rate of metabolism under most conditions, the
Anhinga generally dissipates a larger portion of its metabolic heat

production (Hj by evaporation (H ) than does the cormorant (Figs. 13,
14).


Figure 11. Relationship between log evaporative water loss and ambient temperature as measured
for 4 Anhingas both with and without incident radiation


LOG EWL (mg H20/ min)
Anhinga anhinga
20-
25 30 35 40
ENVIRONMENTAL TEMPERATURE (C)
100
10
LOG EWL (mg H20/min)


Figure 12. Relationship between log evaporative water loss and ambient temperature as measured
for 5 Double-crested Cormorants both with and without incident radiation


LOG EWL (mg H20/min)
Phalacrocorax auritus
y
/'

INCIDENT
RADIATION 700 w/m2
*
O
WITHOUT
INCIDENT RADIATION
1.0-

INCIDENT
RADIATION = 450 w/m2
i i 1 1 1 1 1 1 1 1
20 25 30 35 40 45
ENVIRONMENTAL TEMPERATURE (C)
LOG EWL (mg H20/min)


Figure 13. Relationship between the ratio of He/Hm and ambient
temperature as measured for 4 Anhingas both with
and without incident radiation


He / Hm
60
I.O-i
0.5-
Anhinga Anhinga

o.o
1.5
1.0
0.5 i
0.0
2.0
1.5
1.0 -
0.5
INCIDENT RADIATION 700 w/m2


INCIDENT RADIATION 450w/m2


INCIDENT RADIATION 0 w/m2
0.0 J
20
30 40 50
ENVIRONMENTAL TEMPERATURE (C)


Figure 14. Relationship between the ratio of He/Hm and ambient
temperature as measured for 5 Double-crested
Cormorants both with and without incident radiation


He / Hm
62
Phalacrocorax auritus #
INCIDENT RADIATION = 700 w/m2
INCIDENT RADIATION = 450 w/m2
3.0-1 t t 1
1.5-1
1.0-
INCIDENT RADIATION = 0 w/m2
i
T
20 30 40
ENVIRONMENTAL TEMPERATURE (C)


63
Anhingas were not observed to guiar flutter during any metabolic
trial without incident radiation (day or night) even when T = 45C.
a
In cormorants, weak guiar flutter first appeared at ambient tempera
tures of from 35C to 37.5C (daytime) and generally increased in in
tensity as Ta rose. In one instance guiar flutter was observed in a
cormorant at night (Ta = 39C, = 41.8C).
Linder incident light, Anhingas exhibited guiar flutter at ambient
2
temperatures as low as 29C with incident radiation = 450 W/m and as
'low as26.8C wtr iYiCideTTtTadiatfon ~ ?G0~W/m^ Guiar flutter was
observed in all trials in which ambient temperature exceeded 33.5C
(450 W/m^) and 32C (700 W/m^) and increased in intensity as "T rose.
a
Guiar flutter in cormorants exposed to incident radiation first appeared
at T = 33C (incident radiation = 450 W/m^) and at T = 20C (incident
a a
p
radiation = 700 W/m ). Guiar flutter was present in all trials in
which Ta exceeded or equalled 33C (450 W/m^) and 29C (700 W/m^)
and increased in intensity as T rose.
a
Plumage Wetting
Metabolic rates measured in birds with wet plumage, sitting in
16 cm of water, averaged 78-130% higher (Anhingas) and 15% higher
(cormorants) than those measured in dry birds during the day. Body
temperatures measured during exposure to these wet conditions ranged
from 36.8C to 39.8C with an average Tfa of 38.72C 0.77 (SD) in
Anhingas and 39.8C to 42C with an average Tb of 40.56C 0.44 (SD)
in cormorants..
Thermal conductances measured in birds with wet plumage averaged
3
0.0929 cm 02/g-h-C (Anhingas), which is 205% of that measured in dry


64
3
Anhingas during the day, and 0.0t>32 cm O^/g-h^C (cormorants), which
is 144% of that measured in dry cormorants during the day.
During the experimental trials featuring wet subjects, the An
hingas appeared to become more saturated with water, remain saturated
longer, and were not able to stand as far out of the water as were the
cormorants. In fact, two of the Anhingas did not even attempt to stand
out of the water and were completely immersed for the entire two hours
in the chamber. Metabolic rates and thermal conductances of these in
dividuals were higher than those of the individuals that were able to
extend the upper half of the body out of the water.
In the metabolic trials featuring wet plumage, Anhingas reached
the highest metabolic rates recorded under any experimental conditions
during the study, possibly approaching maximal rates of oxygen con
sumption for this species, with V reaching 4.11 times basal meta
bolic rate in one soaked individual exposed to water at 15.8C
(T^ = 37.4C). Cormorant metabolic rates recorded under similar
conditions were no higher than those recorded at lower temperatures
in dry individuals.


CHAPTER FIVE
DISCUSSIONFIELD OBSERVATIONS
Anhingas, Double-crested Cormorants, and Flightless Cormorants
are highly effective predators, often capturing three to five fish in
a 5 to 10 minute period. As a result these species devote the
majority of their time to activities other than foraging. All three
species, therefore, can be considered as "time minimizers" with re
spect to feeding strategy (Schoener 1971). Their reduced buoyancy,
due to the water absorbing plumage, may improve their efficiency as
underwater predators, at the extreme permitting the Anhinga to remain
relatively motionless under water. However, the temporary increase
in thermal conductance that results from having a layer of water
adjacent the skin must subject them to thermoregulatory stress,
especially in Anhinga (Mahoney in press), a predicament similar to
that suffered by land birds exposed to heavy rains (Kennedy 1970).
This may explain why Anhingas spend so little time in the water,
especially during cool weather.
Considering the high rates of heat loss that must occur in indi
viduals with saturated plumage, it would be energetically advantageous
for Anhingas to dry as quickly as possible after foraging, thus re
establishing a layer of air next to the skin. Spread-winged behavior
may be an adaptation to this end. Such a function is suggested by
65


66
the appearance of this behavior in wet birds on overcast days, and by
their fan drying behavior, which almost certainly functions to dry
the plumage rapidly when sunlight is unavailable. Anhingas often ex
pose the skin during spread-winged behavior by erecting wing and back
feathers, especially while wet. Rapid drying should be facilitated by
a combination of feather erection, the black feather pigmentation, and
exposure of the maximum surface area to the sun via wing extension and
orientation. Although the remiges may not absorb water to the extent
of the breast feathers studied by Casler (1973) and Rijke (1968), the
wing coverts may be wettable, and it may be these which the bird is
attempting to dry by fanning when the sky is overcast. Although the
wings appear water laden when birds leave the water, the contention
that Anhingas must dry their plumage before they can fly (McAtee and
Stoddard 1945) is not supported by personal observations of birds fly
ing, albeit weakly, from the water's surface. This suggests that any
wing drying function serves primarily as an adaptation to conserve
metabolic heat.
Although a wing drying function for spread-winged behavior is
supported by these observations, other data suggest that spread-winged
behavior in Anhinga also has an overlapping thermoregulatory function.
(1) Dry Anhingas display this behavior more often than do wet
birds.
(2) The frequency of this behavior is inversely correlated with
Tfl and may comprise as much as 88% of the behaviors observed with Tg
is less than 10C (Fig. 1). In contrast, the frequency is only 14% at
temperatures above 30C.


67
(3) Anhingas are black on the back and wings. Darkly pigmented
plumage may absorb more solar radiation than light (Hamilton and Hepp-
ner 1967 a&b, Lustick 1969) depending on the wind speed (Walsberg
et al. 1978) and the angle of incidence (Lustick 1980). That Anhingas
normally gain heat from insolation is obvious because guiar flutter
appeared in free-living birds when Ta was between 25.0 and 27.5C and
the intensity of solar radiation was moderate but did not appear in
my captive birds even when chamber temperatures exceeded 37C, unless
a radiant load was present.
(4) Most birds oriented away from the sun (perpendicular to the
incident radiation) during spread-winged behavior but faced the sun
during heat stress (guiar flutter). Orienting perpendicular to
incident radiation would enable Anhingas to take full advantage of
the heat absorbing qualities of the black plumage by increasing the
surface area exposed to the sun and by optimizing the angle of inci
dence (Lustick 1980). Facing the sun would reduce the amount of
surface exposed to it, as demonstrated for Herring Gulls, Larus
argentatus (Lustick et al. 1978), and reduce the angle of incidence,
making the birds effectively white with respect to the absorption
of radiant energy (Lustick 1980).
(5) Anhingas exhibit spread-winged behavior before entering the
water. This may enable them to store heat passively to offset the
high thermal conductance associated with wetting.
(6) During spread-winged behavior Anhingas expose the darkly
pigmented skin of the subalular apterium to the sun. The apterium
is the only darkly pigmented skin on the Anhinga's body and may play
a role in enhancing insolation absorption.


68
The spread-winged behavior of Anhingas is not a mechanism for
"dumping heat" since it never accompanies guiar flutter. In addition,
spread-winged sunning in Anhingas is not simply an adaptation to re
align the feathers after soaring as proposed for vultures and other
soaring birds by Houston (1980). Although Anhingas occasionally soar,
they maintain spread-winged postures for much longer periods of time
than would be required for this purpose, and the appearance of this
behavior is not in any way associated with soaring.
The plumage of Double-crested Cormorants, in contrast with that
of Anhingas, is only partially wettable, as only the distal portions
absorb water. They consequently maintain an air layer next to the
skin while foraging (Casler 1973). This difference in feather struc
ture affords cormorants greater insulation while immersed and while
drying and may explain why cormorants apparently spend more time in
the water than do Anhingas.
Spread-winged behavior in Double-crested Cormorants appears to
function primarily, or solely, in wing drying and not to supplement
metabolism, as
(1) Cormorants only displayed spread-winged behavior when wet,
often accompanied by fanning behavior
(2) Spread-winged behavior comprised too small a percentage of
behavioral observations to allow sufficient time for any significant
supplementation of metabolism (2.3% overall and 4.6% maximum under any
environmental conditions as compared to 30.9% overall and 88% when
sunny and cool for Anhingas)
(3) Double-crested Cormorants did not flatten out to the sun as
Anhingas did but stood erect regardless of sun position


69
(4) Double-crested Cormorants oriented more strongly to the wind
than to the sun.
As with Anhingas, the spread-winged behavior of Double-crested
Cormorants is not a mechanism for "dumping heat" as it was never ac
companied by guiar flutter. The delta-winged posture of these cormor
ants may, however, function in heat dissipation (bringing arterial
blood from the brachial artery close to an exposed, often shaded,
bodily surface), as it was only observed in conjunction with guiar
flutter, and only at high ambient temperatures and solar intensities.
An alternate, but (in my opinion) uni ikely, hypothesis for the function
of spread-winged behavior in cormorants is use as a social signal to
indicate foraging success (Jones 1978).
There are similarities in the use of spread-winged behavior for
both cormorant species (Flightless and Double-crested) as compared with
Anhingas. All three orient nonrandomly with respect to the sun during
spread-winged behavior, with the primary skew in the data being in favor
of orienting with back to the sun. However, the tendency is much
stronger in Anhingas. In fact, the orientation during spread-winged
behavior of Flightless Cormorants was not significantly different from
their orientation during other behaviors. None of the three apparently
use spread-winged behavior for heat dissipation (Flightless Cormorants
do not display spread-winged postures in conjunction with guiar flut
ter either). The drooped-winged posture of Flightless Cormorants (as
with the delta-winged posture of Double-crested Cormorants) may, how
ever, serve this purpose.
There are, however, important differences in the use and function
of this behavior in the Flightless Cormorant (as with the Double-crested


70
Cormorant) in comparison to Anhingas. Because both species possess
wettable plumage, it was thought that perhaps Flightless Cormorants
might share the Anhinga's tendencies towards the supplementation of
metabolism, as these cormorants must forage in the cool waters of the
Humboldt Current (the nearshore surface temperature of which averaged
20.9C during my stay at Tagus Cove). This does not appear to be
the case as Flightless Cormorants
(1) do not orient strongly to the sun,
(2) do not exhibit spread-winged postures while dry,
(3) do not expose a maximum of surface area to the sun by flat
tening out when the sun is high,
(4) actually suffered a slight decline in body temperature
during spread-winged behavior during my measurements.
In addition to these observations, Flightless Cormorants observed
during the course of this study were frequently observed to be guiar
fluttering because, although ambient temperature never exceeded 30C,
black-body temperature frequently exceeded 50C. Frequently this
guiar flutter continued until well after the sun had set, indicating
that perhaps these Flightless Cormorants have high rates of metabolism
to deal with the cool waters in which they forage. These observations
suggest that the heat lost during foraging might be a welcome relief,
as opposed to an energetic stress. It therefore appears that spread
winged behavior in Flightless Cormorants (as with Double-crested
Cormorants) is primarly a mechanism for wing drying. Why this might
be necessary remains a mystery. It may be for energetic reasons (to
reduce heat loss by rapidly drying wet plumage) or even a vestigial
behavior. However, it is not a mechanism for gaining heat from the sun.


CHAPTER SIX
DISCUSSIONLABORATORY RESULTS
Both Anhingas and Double-crested Cormorants show a daily variation
in energetics. These diurnal species reduce their basal rates, body
temperatures, and thermal conductances during the night. However,
neither species displays the 25% reduction in basal metabolic rate
expected from Aschoff and Pohl (1970) or the 50% reduction in thermal
conductance expected from Aschoff (1981).
Anhingas are capable of saving significant amounts of energy by
lowering their body temperature and thermal conductance at night when
exposed to temperatures below thermoneutrality. This adaptation en
ables them to extend the thermoneutral zone 3.5C lower at night than
during the day. Total energy expenditure for thermoregulation can be
calculated from the equation M = C(Tb Tg), where M = metabolic rate
and C = thermal conductance (McNab 1974). Using daytime values for
3
T^ and C and assuming 1 cm Or, = 4.8 cal, total energy expenditure for
a 1.04 kg Anhinga would amount to 188.74 kcal/day used for thermo
regulation at 5C. Using nighttime values for C and T^, and assum
ing 12 hours of night, total daily energy expenditure for thermoregu
lation at 5C would amount to 166.33 kcal/day, a 22.41 kcal savings.
Therefore, in addition to extending the thermoneutral zone, reducing
thermal conductance and body temperature at night enables an Anhinga
to save 12% of the energy it would spend for thermoregulation at 5C.
The slight reduction in nocturnal basal metabolic rate represents a
71


72
savings of 8% of the daily energy expenditure at ambient temperatures
within thermoneutrality (assuming 12 hrs of night). By saving energy
in this manner Anhingas can compensate somewhat for their high thermal
conductances, high rates of heat loss during aquatic behavior, and
limited thermoneutral zone. A reduction in daily energy needs trans
lates into reduced food needs, which means that less time need be
spent in the water and/or drying out. Further reduction of daily en
ergy requirements in Anhingas is accomplished by absorbing insolation
using the spread-winged posture, to be discussed below.
Like Anhingas, Double-crested Cormorants also extend their thermo
neutral zone 3.5C lower at night than during the day. Using daytime
values for C and a 1.33 kg cormorant would expend 204.1 kcal/day
for thermoregulation at 5C. Using nighttime values for C and T^,
and assuming 12 hours of night, total daily energy expenditure for
thermoregulation would amount to 188.34 kcal/day at 5C, a 15.76 kcal
savings. Therefore, in addition to extending the thermoneutral zone,
reducing thermal conductance and body temperature at night enables a
cormorant to save 8% of the energy it would spend for thermoregulation
at 5C. The reduction in basal metabolic rate observed at night in
cormorants represents a savings of 13% of the daily energy expenditure
at ambient temperatures within thermoneutrality (assuming 12 hours of
night). Double-crested Cormorants have high thermal conductances, as
well as high basal rates. Saving energy in this fashion may help the
cormorant to balance its energy budget. Because Double-crested Cormor
ants apparently do not suffer the same degree of thermoregulatory
stress during wetting as Anhingas, theymay not be under as intense


73
selection to evolve energy conserving adaptations such as facilitated
insolation absorption.
The use of insolation to reduce the metabolic costs of endothermy
contrasts greatly between the two species. Double-crested Cormorants
apparently do not use the spread-winged posture to absorb insolation
and reduce their metabolic costs. Their spread-winged behavior prob
ably functions solely in drying the plumage, as has been proposed for
other cormorant species (Rijke 1968, Siegfried et al. 1975). In con
trast to Anhingas, the cormorant's feather structure which maintains
an insulative air layer next to the skin while immersed (easier 1973),
in combination with their high rates of metabolism which permits them
to have a broader thermoneutral zone than Anhingas, may have enabled
Double-crested Cormorants to fully invade the temperate zone. The
feather structure and accompanying suite of morphological/physiological
adaptations of Double-crested Cormorants is consistent with their for
aging strategy, one of active pursuit (Owre 1967).
In contrast to cormorants, both metabolic data and field observa
tions (Hennemann 1982) suggest that Anhingas use spread-winged behavior
to absorb insolation, which hastens the drying of their body plumage,
supplements their low metabolic rates, reduces their high thermal con
ductances during cold weather, replaces body heat lost through convec
tion and evaporation during and after foraging, and allows them to
extend the region of thermoneutrality. The Anhingas' world-wide re
striction to areas with tropical or subtropical climates agrees with
the data on basal metabolic rate and thermal conductance in suggesting
that Anhingas are poorly adapted to and may be energetically stressed
during cold weather. In northern Florida Anhingas are near their


74
northernmost winter limit of their geographic distribution (Palmer
1962). The rather high lower limit of thermoneutrality of Anhingas
means that Anhingas face ambient temperatures below thermoneutrality
for much of the north Florida winter. Another source of potential
energetic stress acting on Anhingas is their wettable plumage. The
Anhingas1 plumage, which allows water to fully penetrate air spaces
next to the skin (Casler 1973), may enhance prey capture but may also
be primarily responsible for the inability of Anhinga to tolerate frigid
weather and fully invade the temperate zone. Due to the many sources
of energetic stress operating on the Anhinga in the temperate zone, the
ability to absorb insolation may be'an essential adaptation to balance
the energy budget of this species in many parts of its range.
Although spread-winged behavior is superficially identical in both
Anhingas and Double-crested Cormorants, the underlying physiological
consequences are quite different. Anhingas may have evolved a thermo
regulatory function after first evolving spread-winged behavior to
facilitate wing drying. The development of its distinctive form of
nearly sedentary stalking fishing behavior (Owre 1967) may have required
the wetting of its plumage to reduce buoyancy, thereby increasing heat
loss and requiring the evolution of a behavioral mechanism to supplement
metabolism. This represents the first documentation of a thermoregu
latory/energetic function for a spread-winged posture and warrants
further investigation of similar behaviors in the many other species
that exhibit them.
At high ambient temperatures, energy absorption by Anhingas may
be a source of thermoregulatory stress rather than advantage. Anhingas
may reduce the risks of heat stress by


75
(1) having low rates of metabolism;
(2) having high thermal conductances;
(3) having a high capacity for evaporative heat loss;
(4) having a capacity for heat storage;
(5) behavioral changes in orientation to the sun. Anhingas primar
ily face the sun during guiar flutter and away from the sun (perpendicu
lar to the incident radiation) during spread-winged sunning (Hennemann
1982). These orientation adjustments enable Anhingas to become ef
fectively white or black as the situation requires (Lustick 1980).
(6) not engaging in spread-winged behavior.
The higher basal metabolic rates and lower evaporative cooling
capacities of cormorants make them less well adapted to the hot, humid,
subtropical summers of Florida. They began guiar fluttering at lower
temperatures than Anhingas, were less capable of dissipating as great
a fraction of their metabolically produced heat through evaporative
means than were Anhingas, and their ratio of H./H exceeded 1.0 at lower
e m
ambient temperatures than in Anhingas.
Incident radiation elevated rates of evaporative water loss in
both species and lowered the ambient temperature at which the ratio of
He/Hm exceeded 1.0. Were it not for the virtually limitless supply of
water available for evaporative cooling, these elevated rates of
evaporative water loss might be a serious source of stress for these
species in hot, sunny environments such as Florida.
In this respect, the wettable plumage of these species, which may
be a source of thermoregulatory stress in the winter, may be an advan
tage in rapid heat dumping in the summer. My examination of the ef
fects of wetting on the thermoregulation of these species is not


76
analogous to field conditions (there are few, if any, situations in
which an Anhinga would remain in the water for two hours, Hennemann
1982). In addition, the great variability in the degree to which dif
ferent individuals were exposed to plumage wetting due to the varia
tion in individual behavior and in the abilities of different indi
viduals and species to stand partially out of the water was another
source of error in making any kind of quantitative assessment of the
effects of plumage wetting on thermoregulation. However, the elevated
metabolic rates and thermal conductances, and reduced body temperatures
observed during wetting, are probably qualitatively representative of
the actual effects of plumage wetting on these two species. The con
tention that Double-crested Cormorants are less stressed by wetting
than are Anhingas is supported by (in addition to the data presented
in this study) behavioral observations that reveal that Anhingas spend
minimal amounts of time in the water, especially at low ambient tempera
tures (Hennemann 1982), while cormorants show no such aversion
(Hennemann, personal observations).


CHAPTER SEVEN
ENERGETICS, BEHAVIOR, AND ZOOGEOGRAPHY
The geographic limits to the distribution of vertebrates are usu
ally difficult to explain. Species may be limited by a number of
factors including physiological and ecological inabilities to cope
with environmental extremes and limitations in temperature, moisture,
resource availability, or a combination of these. With this in mind,
the northern limits of distribution'during winter of Anhingas and
Double-crested Cormorants were examined to delineate and compare the
physiological factors operating on these two species at or near their
geographic limits. To this end, the December distribution of Anhinqa
anhinga in the U.S. as determined from Christmas Audubon Society bird
censuses (published in the July issues of American Birds for 1977-1981)
is presented in Figure 15. The December distribution of Phalacrocorax
auritus in northeastern North America, as determined from Audubon
Society Christmas censuses from 1977-1981, is presented in Figure 16.
The northern limit to the distribution of Anhingas in eastern
North America does not appear to depend on ecological factors such as
the availability of food or open water but rather on two interacting
parameters of the physical environment, ambient temperature and the
availability of insolation. Specifically, in December Anhingas can be
found in localities where the mean monthly temperature (for December)
is 10C, provided that there is also 160 or more hours of sunshine
available per month. If the mean hours of monthly sunshine for
77


Figure 15. Distribution of Anhingas in the U.S
in December




Figure 16. Distribution of Double-crested Cormorants in northeastern North America in
December




82
December are less than 160 hours per month, then Anhingas may be found
between the 10C and 12.5C mean monthly temperature isotherms (closer
to the 12.5C line) such as in southern Mississippi, or they may drop
out entirely, as is the case in south Alabama near Mobile, and in north
western Florida near Pensacola. Anhingas extend north of the 10C
isotherm in only two areas: the eastern coasts of the Carolinas as
far north as Wilmington, North Carolina, and in northwestern Louisiana
near Shreveport, and southwestern Arkansas as far north as Texarkana,
Arkansas. Not surprisingly, these areas receive an average of better
than 175 hours of sunshine in December (Wilmington--l78, Shreveport--
177, Texarkana--no available data)J'
Apparently, Anhingas can balance their energy budgets in two ways:
(1) by living in areas subject to moderate temperatures or (2) by
living at less moderate temperatures in areas in which there is ample
sunshine to supplement metabolic heat production. A series of energy
budgets constructed using my behavioral and energetic data is pre
sented in Tables 11-13. These energy budgets illustrate the energetic
costs for thermoregulation for a 1.04 kg Anhinga assuming that the
average temperature is 10C and (1) sun is available and the Anhinga
is exhibiting behavioral frequencies appropriate for a sunny period
at temperatures at or below 15C (Table 11), (2) sun is not available
(skies are overcast) and the Anhinga is behaving appropriately for a
cloudy period in which temperatures are at or below 15C (Table 12, and
(3) the sun is not available but the Anhinga is exhibiting behavioral
frequencies appropriate for a sunny day, not a cloudy one (Table 13).
^Climatic data were obtained from the Climatic Atlas of the United
States, 1968 edition, a publication of the U.S. Department of Commerce


Table 11. Daily energy budget for thermoregulation (Anhinga)
assuming
1) Daytime Ta averages 15C (10 hours of day)
2) Nighttime Ta averages 6.5C (14 hours of night)
3) Water temperature averages 10C
4) Sun is available
5) Anhinga
behavior is
appropriate
for a sunny day
Behavior
Frequency9
(%)
C b
m
(cnrO^/ghr0
Tb
'0 (c)

(C)
MC
(cm302/g hr)
Madj
Costs6
(kcal/day)
S-W Sunning
(Wet)^
20.4
0.0605
39.4
24.4
1.48
3,140
15.07
S-W Sunning
(Dry)9
43.3
0.0281
40.0
25.0
0.70
3,516
16.88
Perching*1
20.2
0.0454
39.7
24.7
1.12
2,353
11.30
Swimming
11.1
0.0929
38.7
28.7
2.67
3,082
14.79
Total (Day)
57.76
Night
100.0
0.0364
38.0
31.5
1.15
16,690
80.13
Total (overall)
137.89


aBehavioral Frequency as a percent of total observations (see text Chapter Two)
bMinimal thermal conductance; Cm for S-W Sunning (Wet) was assumed to be the average of Cm as measured
for dry birds exposed to insolation and C as measured for wet birds
cMetabolic rate as calcuated from the equation M = (T^ Ta) (McNab 1974)
^Adjusted metabolic rate calculated by multiplying M times the proportion of time devoted to that
behavior times 10 hours (day) or 14 hours (night) times Mass, expressed in cm^O^/lO hr or cnr^/M hr
eassumes 1 cm302 = 4.8 cal
^Spread-winged sunning with wet plumage
^Spread-winged sunning with dry plumage
^Perching includes all behaviors except swimming and sunning, the assumption being that all such
behaviors should have similar values for C
m
CO
-p


Table 12. Daily energy budget for thermoregulation (Anhinga)
assuming
1) Daytime Ta averages 15C (10 hours of day)
2) Nighttime TQ averages 6.5C (14 hours of night)
3) Water temperature averages 10C
4) Skies are overcast
5) Anhinga
behavior is
appropriate for
a cloudy day
Behavior
Frequency
(%)
^m
(cm302/ghrC)
Tb
(c)
VTa
(C)
M
Madj
Costs
(kcal/day)
S-W Sunning (Wet)a
6.1
0.0692b
39.2
24.2
1.67
1,077
5.17
Perching
89.0
0.0454
39.7
24.7
1.12
10,367
49.76
Swimming
4.9
0.0929
38.7
28.7
2.67
1 ,360
6.53
Total (day)
61.46
Night
100.0
0.0364
38.0
31.5
1 .15
16,690
80.13
Total (overall)
141.59
aincludes fan drying
^assumed to be the average of C as measured for dry birds without insolation and C as measured for
wet birds m


Table 13. Daily energy budget for thermoregulation (Anhinga)
assuming
1) Daytime Ta averages 15C (10 hours)
2) Nighttime Ta averages 6.5C (14 hours)
3) Water temperature averages 10C
4) Skies are overcast
5) Anhinga behavior is appropriate for a sunny day, not cloudy
Behavior
Frequency
(%)
^m
(cm302/g hC)
Tb
(c)

(C)
M
Madj
Costs
(kcal/day)
S-W Sunning (Wet)
20.4
0.0692
39.2
24.2
1.67
3,543
17.01
S-W Sunning (Dry)
48.3
0.0454
39.7
24.7
1.12
5,626
27.00
Perching
20.2
0.0454
39.7
24.7
1.12
2,353
11.29
Swimming
11.1
0.0929
38.7
28.7
2.67
3,082
14.79
Total (day)
70.09
Night
100.0
0.0364
38.0
31.5
1.15
16,690
80.13
Total (overall)
150.22


87
These time and energy budgets suggest that Anhingas often delay
feeding on cool, cloudy days, compensating for this by feeding heavily
during sunny periods. Even with reduced time in the water on cloudy
days, Anhingas still expend 6.4% more energy for thermoregulation than
they would on a sunny day in which they spend significantly more time
in the water. Table 13 indicates that if sunny periods were scarce and
Anhingas were forced to forage as much on cloudy days as they do on
sunny days, their daytime costs for thermoregulation would rise by
21%, possibly causing them to exceed the limits of their energy budget.
This offers an explanation for the apparent dependency of Anhingas on
either very moderate temperatures, or frequent sun, or both, as sug
gested by their distribution.
The distribution of Double-crested Cormorants, on the other hand,
apparently is not limited by physical factors such as T or insolation,
cl
as the coolest areas in which they are found are also the regions with
the fewest number of sunny hours per month (upstate New York, southern
Quebec with fewer than 80 hours of sunshine in December as compared
to coastal regions such as near Portland, Maine, which receives over
120 hours of sunshine in December and more moderate temperatures). A
more likely explanation for the northern limit of the distribution of
this species is an ecological factor, such as the availability of
open water.
Because Double-crested Cormorants do not use the sun to supplement
metabolism, I constructed energy budgets for them using pooled behav
ioral frequency data from both sunny and cloudy periods at temperatures
at or below 15C. Table 14 illustrates the thermoregulatory costs for
a 1.33 kg Double-crested Cormorant assuming that ambient temperatures


Table 14. Daily energy budget for thermoregulation (Double-crested Cormorant)
assuming
1) Daytime Ta averages 15C (10 hours of day)
2) Nighttime Ta averages 6.5C (14 hours of night)
3) Water temperature averages 10C
Behavior
Frequency
(%)
3 Cm
(cmJ02/g hC)
Tb
(C)

(c)
M
Hadj
Costs
(kcal/day)
Spread-winged
2.6
0.0450
40.9
25.9
1.17
405
1.95
Perching
69.9
0.0365
41 .0
26.0
0.95
8,832
42.39
Swimming
27.5
0.0532
40.6
30.6
1 .63
5,962
28.62
Total (day)
72.96
Night
100.0
0.0320
39.7
32.2
1 .03
19,179
92.06
Total (overall)
165.02


89
average 10C, to compare costs with the Anhinga at this temperature.
Table 15 presents the costs faced by these cormorants near their
northernmost distributional limit. Although Double-crested Cormorants
expend more energy than do Anhingas for thermoregulation at 10C av
erage temperature, they expend less energy on a per gram basis (per
unit body mass). Because their plumage is only partially wettable,
cormorants can, and do, spend more time in the water than do Anhingas
at these ambient temperatures. They, therefore, are capable of cap
turing more prey (presumably) than their completely wettable relatives,
at lower expense (as is suggested by the data on the effects of wetting
on metabolism). This ability to forage freely on cool, cloudy days
offers an explanation for the ability of the Double-crested Cormorant
to live at latitudes far north of the Anhinga's northern limit and at
ambient temperatures far lower than the Anhinga can tolerate.
The energy budgets confirm what the field and laboratory data im
plied. The dependency of the Anhinga on a "stalking" foraging stragegy
featuring totally wettable plumage tied to low rates of metabolism,
high thermal conductances, and a dependency on solar assistance restricts
Anhingas to subtropical climates with warm winters and ample sunshine,
while the "chasing" strategy of the Double-crested Cormorant featuring
partially wettable plumage tied to high metabolic rates, high con
ductances, and no special adaptation for insolation absorption allows
these cormorants to extend well into the temperate zone.


Table 15. Daily energy budget for thermoregulation (Double-crested Cormorant)
assuming
1) Daytime Ta averages 0C (10 hours of day)
2) Nighttime Ta averages -8.6C (14 hours of night)
3) Water temperature averages 1C
Behavior
Frequency
(%)
r
m
(cm302/g hC)
Tb
(C)

(C)
M
Madj
Costs
(kcal/day)
Spread-winged
2.6
0.0450
40.9
40.9
1.84
636
3.05
Perching
69.9
0.0365
41.0
41 .0
1.50
13,945
66.94
Swimming
27.5
0.0320
40.6
39.6
2.11
7,717
37.04
Total (day)
107.03
Night
100.0
0.0320
39.7
48.3
1.55
28,861
138.53
Total (overall)
245.56


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Full Text



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