INTEGR. COMP. BIOL.

, 45:219230 (2005)

George A. Bartholomews Contributions to Integrative and Comparative Biology1

WILLIAM R. DAWSON2
Museum of Zoology, The University of Michigan, Ann Arbor, Michigan 48109-1079

INTRODUCTION
I want to note my pleasure at being a participant in
a symposium that both honors George Bartholomew
(Bart to his students and other friends) and commemorates the tenth anniversary of the George A. Bartholomew Award. This award, which recognizes gifted
young investigators in integrative and comparative biology, is a tangible indication of the esteem in which
Bart is held by his colleagues in the Division of Comparative Physiology and Biochemistry of The Society
of Integrative and Comparative Biology (SICB). I have
been asked to comment on Barts contributions to integrative and comparative biology. I have arbitrarily
divided his accomplishments in this regard into three
categories: definition of precepts, original research,
and education and service. I shall comment on these
in order.

themselves into categories and then pretend that these

categories exist in the living systems under investigation (Bartholomew, 1958). Thus he has emphasized
that animals are indivisible functionally and that his
field of physiology is not an isolatable organismal
component, being inextricably linked with morphology and behavior. He pursued this view of a unified
biology further in an eloquent statement concerning
modern natural history (Bartholomew, 1986, p. 329)
that has broad implications for integrative and comparative biology: Because of its focus on organisms
natural history is in a unique position to supply questions and integrating links among disciplines. Studies
at lower levels will delineate the machinery of structural units, and the complex systems into which these
units have been assembled through evolutionary time
will be worked out by research at the intermediate and
higher levels of biological integration. Biology is indivisible; biologists should be undivided. Bart (1964,
p. 8) had provided earlier justification for an inclusive
view of biology through recognition of this hierarchy
of biological explanations: There is a familiar solution to this problem [parochialism based on differences
in approach by various biologists], widely recognized
but sometimes difficult to accept emotionally. This is
the idea that there are a number of levels of biological
organization and that each level offers unique problems and insights, and further, that each level finds its
explanation of mechanism in the levels below, and its
significance in the levels above.
As is evident from his contribution to this symposium (Bartholomew, 2005), George Bartholomew has
also made it a rule to explore the evolutionary implications of his research wherever feasible. One of the
foundations of this practice has been his appreciation

DEFINITION OF PRECEPTS
George Bartholomew in his valedictory for this
symposium (Bartholomew, 2005) discusses creativity
and the precepts that have guided his scholarly activities over his career. Through his adherence to these
precepts, he has been able to exert a powerful effect
on the post-World War II development of ecologically
oriented physiology. His holistic view of biology continues to inspire in this age of disciplinary fractionation and to convey an important message. He has recognized that biology is a continuum, but that biologists
because of their human limitations tend to divide
1 From the Symposium Integrative Biology: A Symposium Honoring George A. Bartholomew presented at the Annual Meeting of
the Society for Integrative and Comparative Biology, 59 January
2004, at New Orleans, Louisiana.
2 E-mail: wrdawson@umich.edu

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SYNOPSIS. The Bartholomew Award has now completed a decade of recognizing outstanding young investigators in comparative physiology and biochemistry or in related fields of functional and integrative biology.
It honors Professor George A. Bartholomew (Bart to his many students and other friends), whose research
contributions continue to be important in shaping these fields. Barts influence reflects a steadfast adherence
to a set of basic precepts: the inherent unity of biology; the need for an evolutionary perspective in functional
studies; the value of modern natural history in guiding research investigations; the focus on the organism
and its function in nature, even in highly reductionist studies; the importance of biological variability within
and between species; and the crucial interactions of physiology and behavior in allowing animals to deal
with environmental challenges. Were he to have done nothing else in his career, he would remain an important figure in the fields with which the Society of Integrative and Comparative Biologys (SICB) Division
of Comparative Physiology and Biochemistry is concerned. However, his influence is also felt through his
inspirational performance as an undergraduate teacher, his skill and wisdom as a graduate mentor, his
many services to the University of California, his insightful contributions to scientific committees and policy
boards at the national level, and his presidency of the American Society of Zoologists (now SICB). This
symposium offers the opportunity for honoring Bart for all his accomplishments and fine personal qualities,
while illustrating the contributions of the impressive set of younger investigators who are recipients of the
George A. Bartholomew Award.

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WILLIAM R. DAWSON
distinction between the two (organism and environment), he has cautioned that this should only be done
if they are never treated separately.
A further precept arising from the views summarized in Barts contribution to this symposium (Bartholomew, 2005) emphasizes interactions between
functional capacities and behavior of organisms, interactions that are frequently critical for survival in difficult physical conditions (Bartholomew, 1964,
1966a). He has noted that such interactions are especially important for terrestrial animals due to the physical complexity of their environments. These animals
because of their mobility and behavioral capacities can
actively seek out and use those environmental situations that best allow their morphological and physiological abilities to function adequately for survival and
reproduction. Most of these species are less than 1%
the size of man and his domestic animals, and their
environment may include shaded or sunny areas, tunnels, cracks, crevices, burrows, thick underbrush, holes
in logs, and/or nests, making gross climatic indices
often irrelevant (Bartholomew, 1964). These and preceding considerations have led him to expend substantial research effort in the field as well as the laboratory.
RESEARCH
George Bartholomew is a true comparative biologist
and the subjects of his studies include insects, amphibians (Bucher et al., 1982; Ryan et al., 1983), reptiles, birds, and mammals, as well as the plant Philodendron selloum (Seymour et al., 1983, 1984). (Many
of the studies arising from his work on insects and
amniotic vertebrates are cited below.) He has consistently operated at the interfaces of comparative physiology, ecology, and behavior. Pursuit of appropriate
research animals and investigation of interesting problems have led him to work in North and Central America; Australia; Europe; Africa; Antarctica; and a number of islands including the Pribiloffs, Midway, and
New Guinea. The following comments do not comprise a comprehensive review of his total research activities (for example, in the interests of producing an
essay of manageable length, I have omitted mention
of several primarily behavioral studies and have not
fully indicated every publication pertaining to some of
the topics that are considered below), but I have tried
through the references that are included to provide
some sense of the scope of his investigations, creativity and, in a number of instances, truly pioneering efforts.
The studies by Bart and his collaborators are noteworthy for their broad perspective. His initial research,
which has been cited over at least 50 years, concerned
behavior of double-crested and Brandt cormorants,
Phalacrocorax auritus and P. penicillatus, on San
Francisco Bay, CA (Bartholomew, 1942, 1943a, b).
Following a three-year interruption due to World War
II, Bart resumed a publication career that would extend
over the next four decades and be marked by a number
of firsts and technical innovations. His doctoral the-

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of the evolutionary implications of biological variability. Differences between individuals are the raw
materials for evolutionary change and for the evolution
of adaptations, yet of course most physiologists treat
these differences as noise that is to be filtered out.
From the standpoint of physiological ecology, the traditional emphasis of physiologists on central tendencies rather than on variance has some unhappy consequences. Variation is not just noise; it is also the
stuff of evolution and a central attribute of living systems. . . . The physiological differences between individuals in the same species or population, and also the
patterns of variation in different groups, must not be
ignored (Bartholomew, 1987, pp. 32-33). In this connection, he has emphasized (Bartholomew, 1987, p.
33) the value of research efforts that can deal critically
with intraspecific comparisons. Rather than just comparing different species, one should adopt some of the
formats developed for interspecific comparisons in an
attempt to compare breeding populations within species . . . and individuals within the same breeding population. . . . Such efforts should allow investigators to
approach more closely the dynamics of evolutionary
change, and, in appropriate situations, to integrate the
findings of physiological ecology with those of population genetics.
George Bartholomew has also consistently emphasized the important role of historical factors in shaping
the functional characteristics of contemporary species
([each species] has an evolutionary history, which
means its present configuration has been shaped by
natural selection [Bartholomew, 1982a, p. 231]). He
has noted further that it is important in the analysis of
the adjustments of organisms to their respective environments to understand that selection results in adequacy of performance rather than perfection. He also
has reflected on the fact that despite the long-term
probability of extinction, every organism alive today
is part of an enormously long success story in which
each of its ancestors has been sufficiently well adapted
to its physical and biotic environments to mature and
reproduce successfully. It is the intact and functioning
organism on which natural selection operates and such
organisms therefore should be the primary concern of
any biologist who aspires to a broad and integrated
understanding of biology. Consequently, Bart regards
adaptation as so central a theme as to be inseparable
from life itself (Bartholomew, 1987). Undoubtedly, his
views have contributed in a number of respects to the
foundation of the burgeoning field of evolutionary
physiology (Garland and Carter, 1994; Feder et al.,
2000; Kingsolver and Huey, 2003).
Bart has also been careful to observe the principle
first enunciated by Claude Bernard that an organism is
inseparable from its environment (Bartholomew, 1958,
2005). Full knowledge of any species requires familiarity not just with its general surroundings but also its
interaction as a self-maintaining physicochemical system with its special microenvironment (Bartholomew,
1958). Although it is convenient to maintain a verbal

GEORGE BARTHOLOMEWS CONTRIBUTIONS

California (Bartholomew, 1950, 1951), but was concerned primarily with detailed investigations of population biology and reproductive behavior. Major studies were conducted on the social and reproductive behavior of the elephant seal, Mirounga angustirostris
(Bartholomew, 1952); Alaska fur seal, Callorhinus ursinus (Bartholomew and Hoel, 1953); and California
sea lion, Zalophus californianus (Peterson and Bartholomew, 1967). Some important thermal information
was also obtained. Due to the large size and effective
insulation of these animals noted above, terrestrial
breeding can put them at risk of overheating, even in
a cool climate (see, for example, Bartholomew and
Wilkie, 1956). Bart showed how a predominantly Holarctic species, the California sea lion, could extend its
range to the equatorial Galapagos Islands where warm
temperatures and intense insolation prevail. Breeding
male sea lions persistently maintain terrestrial territories on the coastal islands of California and Baja California, where thermal conditions are ameliorated by
cool upwelling water and a persistent summer overcast. The breeding males in the Galapagos use a behavioral strategy in dealing with the heat challenge
prevailing there. During the daylight hours, they protect themselves from overheating by holding aquatic
territories while remaining immersed in tide pools or
channels, hauling out on land only at night (Bartholomew, 1966a). This seemingly simple strategy, imposes profound effects on a breeding structure that is
exclusively terrestrial in other otariids (sea lions and
fur seals), but it is an excellent example of a behavioral
solution to a primarily physiological problem, a theme
that recurs in many of Barts studies.
Early in his career, Bart recognized breeding colonies of seabirds as an excellent resource for thermoregulatory studies. Many of these birds nest in exposed
situations and, at tropical or subtropical latitudes in
particular, they and their eggs and young can be exposed to dangerously intense insolation and high air
temperatures. Commencing with a relatively simple
field study of western gulls, Larus occidentalis (Bartholomew and Dawson, 1952), Bart participated in
several projects concerning the ontogeny of avian thermoregulatory capacity. One involved a comparison of
three species of seabirds that nest concurrently on the
surface of a hot desert island in the Gulf of California,
Mexico, despite major differences in the thermoregulatory proficiency of their hatchlings. The results demonstrated how closely attentive behavior of parents of
each species was attuned to the developmental status
of their chicks (Bartholomew and Dawson, 1954). Further extensive observations of breeding seabirds on
Midway Island, where intense insolation is also a
problem, documented thermally significant behavior of
both parents and young of various ages (Howell and
Bartholomew, 1961a, b, 1962a, b). One lasting image
is of older nestling albatrosses (Diomedea nigripes and
D. immutabilis) with their backs to the sun resting on
their heels and raising their heavily vascularized and
shaded webbed feet off the ground into the cooling

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sis established the thresholds of light intensity necessary for evoking photoperiodic responses in house
sparrows, Passer domesticus (Bartholomew, 1949).
After joining the faculty at UCLA he was the first
physiological ecologist to initiate an organized program of research on the physiology and behavior of
desert birds. This resulted in a number of papers on
the thermoregulatory responses (e.g., Bartholomew
and Cade, 1957a; Bartholomew and Dawson, 1958;
Bartholomew et al., 1962) and on water and electrolyte
metabolism (e.g., Bartholomew and MacMillen, 1960;
Smyth and Bartholomew, 1966) of these animals, culminating in an influential review of water relations
(Bartholomew and Cade, 1963) and a model that accounted for the ability of certain small seed eating
birds to survive without drinking (Bartholomew,
1972). Bart continued to investigate the responses of
birds to heat challenges in several collaborations (Lasiewski and Bartholomew, 1966; Bartholomew et al.,
1968; Lasiewski et al., 1970), obtaining indications in
common poor-wills (Phalaenoptilus nuttallii), doublecrested cormorants, brown pelicans (Pelecanus occidentalis), and mourning doves (Zenaidura macroura)
that gular fluttering, an activity serving to increase
evaporative water loss at high temperatures, occurred
at the resonant frequency of the gular apparatus (Lasiewski and Bartholomew, 1966; Bartholomew et al.,
1968). Such congruence would reduce the energy costs
of this form of cooling.
One particularly important facet of the research on
water and electrolyte balance referred to above documented variation among populations of the savannah
sparrow (Passerculus sandwichensis) in patterns of use
of NaCl solutions or sea water as fluid sources (Cade
and Bartholomew, 1959). Members of two subspecies
resident in salt marshes (P. s. beldingi and P. s. rostratus) increased their drinking with increasing concentration of the solutions provided, whereas three
northern migratory subspecies of savannah sparrow
(P. s. anthinus, P. s. nevadensis, and P. s. brooksi)
decreased theirs, like most other birds. This indicated
marked differences in electrolyte metabolism between
the two groups and the salt marsh residents proved
more proficient in obtaining physiologically useful water from salt and sea water solutions than the migrant
subspecies. A subsequent study (Poulson and Bartholomew, 1962) revealed important physiological differences between representatives of one of the salt marsh
taxa (P. s. beldingi) and those of a migrant form (P.
s. brooksi). The salt marsh residents could tolerate significantly higher serum osmotic pressures and chloride
concentrations and produce substantially more concentrated urine than the migrants. These studies are notable in providing an impressive early example of geographic variation at the intraspecific level in avian
physiological performance.
Another of Barts early research projects dealt with
pinnipeds. This initially produced documentation of
the resurgence of populations of two species of these
animals on islands off Baja California and southern

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WILLIAM R. DAWSON
of this involved a study of the brown pelican, a representative of the Order Pelecaniformes, a group of
particular interest because it includes the largest altricial birds and the only marine birds showing this developmental pattern. Several features of the pelicans
embryonic development differ from those in other altricial species (eyes open at hatching, high energy density of the egg, the large amount of yolk present in
hatchlings, the relation of mass-specific oxygen consumption to embryonic age, and the high total cost of
development up to hatching [121 kJ]). The brown pelican produces a relatively large chick at a relatively
high cost compared to other, smaller altricial species.
The divergence of its pattern of embryonic development from that in these other birds was regarded as
consistent with the hypothesis that avian altriciality is
polyphyletic (Bartholomew and Goldstein, 1984). Data
primarily from the brown pelican and three other species from different orders were used in an analysis
assessing correlates of variation among birds in growth
pattern, gas exchange, and energetics of development
(Bucher and Bartholomew, 1984). These three physiological variables were found to be correlated with
body mass, time, energy, phylogeny, and ecological
niche of the species. The study emphasized the importance of variability as a biological reality and the
deficiencies of confining the analysis embryonic
growth processes entirely to definitions of central tendencies. Bart also collaborated in another study of developmental variation, in this instance dealing with the
Adelie (Pygoscelis adeliae) and emperor penguins
(Aptenodytes forsteri), both of which lay smaller eggs
than large species in other avian orders (Bucher et al.,
1986). The incubation period of the former species
was similar to that predicted on the basis of egg mass,
but that of the emperor penguin was 50% longer. Total
oxygen consumption of developing chicks of the two
species over the incubation period matched values predicted for precocial birds, leading to the conclusion
that these penguins were incorrectly classified as semialtricial.
The role of heterothermy in the biology of certain
birds and mammals that must deal with challenges involving heat, cold, or restriction of food or water, has
long intrigued George Bartholomew. He led or participated in the first successful laboratory studies (Bartholomew et al., 1957, 1962; Howell and Bartholomew, 1959) of the common poor-will, the only bird
known to hibernate. At a time when metabolic level
in a given group of animals was regarded by many as
solely a function of body mass, these studies documented the birds remarkably low standard metabolic
rate as well as its heterothermic capacities and extensive powers of heat defense. Information on dormancy
in swifts and hummingbirds was also obtained (Bartholomew et al., 1957). Bart also collaborated in studies of dormancy in desert rodents (Bartholomew and
Cade, 1957b; Bartholomew and Hudson, 1960; Bartholomew and MacMillen, 1961; Brown and Bartholomew, 1969), analyzing the capacities of pocket mice

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trade winds (Howell and Bartholomew, 1961a). Barts

discerning eye led to a particularly informative account of the behavioral repertoire (orientation away
from the sun, holding its wings away from the body,
elevation of the scapular feathers) used by the masked
booby (Sula dactylatra) while protecting itself and
shading its eggs and young chicks from the intense
solar radiation and high ambient temperatures prevailing in the Galapagos during the breeding season (Bartholomew, 1966b). A later study on another island in
the Gulf of California demonstrated the precise coupling of thermally protective incubation behavior by
adult Heermanns gulls (Larus heermanni) to ambient
conditions (Bartholomew and Dawson, 1979).
Behavior can have a major impact on the thermal
relations of terrestrial as well as aquatic birds. A particularly spectacular example of this is provided by
sociable weavers (Philetairus socius). These weaver
finches, which are the size of house sparrows, construct huge, communal nests in the Kalahari Desert,
which they continuously occupy and maintain. Bart, F.
N. White, and T. R. Howell examined the extent of
thermal protection provided by these structures in
summer and winter (White et al., 1975; Bartholomew
et al., 1976). In the former season, outside air temperatures ranged from 168 to 33.58C, whereas temperatures in occupied nest chambers only varied 78 or 88C.
Winter in the Kalahari often involves sustained nocturnal winds and temperatures that can drop to freezing. At this season, the insulation of the nest in combination with the heat production of roosting birds produced temperatures as high as 378C, 238 above outside
air. Only two sociable weavers occupied a nest chamber in the summer, whereas up to five did so in winter,
leading Bartholomew et al. (1976) to conclude that
changing this number is the principal means by which
these birds maintain chamber temperatures within their
zone of thermal neutrality throughout the year. This
thermostability leads to energy savings of nearly 50%
as compared with estimated costs were the birds to
roost in the open air. This curtailment of thermoregulatory costs appears important both in allowing both
higher population densities of the sociable weaver in
an area of low biological productivity and in the extension of breeding into the cooler parts of the year.
The latter is probably significant in helping this bird
to avoid the heavy reptilian predation on parents, eggs,
and young that can occur in summer. The use of burrows for nesting also can provide thermal protection,
as the study by F. N. White, Bart and J. L. Kinney of
a Spanish nesting colony of the European bee-eater,
Merops apiaster, illustrates (White et al., 1978). However, poor nest sanitation and microbial action on the
accumulated excrement can lead to buildups of NH3
and CO2, which are alleviated by the ventilation resulting from wind and the movement of adults in and
out of the nest tunnel.
Barts later research on avian development involved
participation in studies of growth patterns, gas exchange, and energetics of avian embryos. One example

GEORGE BARTHOLOMEWS CONTRIBUTIONS

limbs and tail (Bartholomew, 1956). The large Australian flying foxes Pteropus poliocephalus and P. scapulatus were found by Bartholomew et al. (1964) to
maintain body temperature in the usual range for mammals at ambient temperatures from 58 to 408C. In cooler surroundings they wrapped their wings about the
body creating an overcoat that allowed skin temperatures to remain as much as 108C above ambient.
At high air temperatures they supplemented their panting by salivation and licking of the wings and chest.
Flapping the wings also contributed to convection over
the body. These behavioral actions contributed to the
ability of the flying foxes roosting in exposed positions
in the tops to trees to tolerate exposure the tropical or
subtropical sun. Behavior also figures prominently in
maintenance of thermal balance in the rock hyrax
(Heterohyrax brucei), a mammal Bart studied during
a sabbatical leave in Kenya. This hyrax has a behavioral repertoire that includes diurnality, basking, restricted periods of surface activity, gregarious habits
within burrows, and huddling, all contributing to effective control of body temperature despite a low metabolic rate and high thermal conductance (Bartholomew and Rainy, 1971). This animal also showed an
unusual physiological response at high temperatures,
increasing evaporative water loss while reducing its
oxygen consumption, the reduction evidently produced
by lowering muscle tonus. Additionally, Bart also participated in a study (Bell et al., 1986) of the energetics
of the leaf-nosed bat (Macrotus californicus), which
involved important behavioral issues. The species is of
special interest due to its being the northernmost representative of a primarily tropical family (Phyllostomidae) and it is does not use torpor in energy conservation. The combined field and laboratory study provided information on thermoregulatory characteristics,
standard and field metabolic rates, and behavior of this
animal. These data supported the conclusion that, rather than relying on special physiological adaptations for
survival, the leaf-nosed bat is successful as a yearround resident in its desert environment through roosting in continuously warm (ca. 298C) geothermal refugia such as caves and mine shafts, and through an
economical method of foraging involving visual prey
detection.
Barts use of the comparative method in desert studies has extended to analysis of reproductive patterns
in several rodents. G. J. Kenagy and he undertook a
long-term investigation of reproductive timing in five
species of rodents (two nocturnal kangaroo rats and
two nocturnal pocket mice in addition to the diurnal
antelope ground squirrel) coexisting in a desert community in the Owens Valley of California. They were
interested in determining whether closely related species differ in reproductive timing as a result of such
things as differences in body size, daily cycle, food
habits, locomotor patterns, and their respective microenvironments. (Kenagy and Bartholomew, 1985).
They found substantial differences among the species,
all of which are long-lived and characterized by rela-

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(Perognathus longimembris), Mohave ground squirrels

(Spermophilus mohavensis), and kangaroo mice (Microdipodops pallidus) for entering torpor and providing the first extensive laboratory observations on the
phenomenon of estivation. This form of dormancy,
which was poorly understood at the time, was explored
further in a review article (Hudson and Bartholomew,
1964).
Bart has extended his interest in mammalian and
avian heterothermy through additional studies involving such species as speckled mouse birds, Colius striatus (Bartholomew and Trost, 1970), pygmy possums,
Cercaertus nanus (Bartholomew and Hudson, 1962),
and two small New Guinea flying foxes: the common
tube-nosed fruit bat, Nyctimene albiventer, and the unstriped tube-nosed bat, Paranyctimene raptor (Bartholomew et al., 1970). The results for the flying foxes
were noteworthy because of the two bats tropical distribution, frugivorous diet, and the fact that a capacity
for heterothermy had not been previously reported for
any members of the Megachiroptera (see Bartholomew
et al., 1964). Additionally, nocturnal torpor involving
body temperatures as low as 26.88C was observed in
two small neotropical birds, the manakins Pipra mentalis and Manacus vitellinus (Bartholomew et al.,
1983). These birds are also frugivorous. Heterothermy
can reduce the energy expenditures of these small animals by more than half. Fruit is not always readily
available in rain forests and Bart and his coauthors
hypothesized that torpor might well occur in other
small tropical songbirds dependent upon such food.
The use of torpor by the Mohave ground squirrel
contrasts sharply with the behavior of the antelope
ground squirrel (Ammospermophilus leucurus) in the
same desert environment. The latter species neither hibernates nor estivates, though it does achieve some
energetic savings by lowering its body temperature to
328338C during winter nights (Chappell and Bartholomew, 1981a). Therefore, it is forced to deal with conditions on the desert surface throughout the year. Its
diurnal habits expose it to severe summer heat, which
it deals with by shuttling between the desert surface
and its burrow. It tolerates hyperthermia when on the
surface, thereby avoiding high rates of evaporative water loss. Periodic retreat to the antelope ground squirrels burrow allows it to dump the heat it has gained,
thereby surviving the stresses of the summer daytime
desert through appropriately timed movements (Bartholomew and Hudson, 1961; Chappell and Bartholomew, 1981a, 1981b).
Thermally relevant behavior is also crucially important for other mammals. In one of the first detailed
studies of thermoregulatory capacity in macropod marsupials (kangaroos, wallabies, etc.), Bart found that the
quokka (Setonix brachyurus) was at least as effective
in its temperature control over an ambient range of
2108 to 448C as eutherian mammals of comparable
size (2.54.0 kg). At high ambient temperatures, this
West Australian species cooled itself by the behavioral
stratagem of spreading saliva over its ventral surface,

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physiological control of body temperature (Bartholomew, 1982b).
The work with the Australian lizards just referred to
featured the first application for reptiles of a concept
developed by F. E. J. Fry (1947) concerning aerobic
metabolic scope (i.e., the difference between the standard and peak rates of oxygen consumption at a particular body temperaturean index of aerobic capacity
for activity) of ectotherms. In an era in which investigators were primarily concerned with standard or
resting metabolic rates, Bart and his collaborators extended their studies to include measurement of the
highest rates of oxygen consumption occurring spontaneously or through stimulation in their lizards (Bartholomew and Tucker, 1963, 1964; Bartholomew et al.,
1965). These studies represented an important first step
in the analysis of the metabolic correlates of activity
and their thermal dependence in reptiles. This has provided part of the stimulus for the development of a
number of studies of the energy cost of locomotion
and the temperature dependence of metabolic scope in
members of this group. Bart has participated in several
of these pertaining to the Galapagos marine iguana
(Bennett et al., 1975; Bartholomew et al., 1976; Vleck
et al., 1981). The studies have examined aerobic and
anaerobic metabolic scope of this species in relation
to temperature. They also have quantified swimming
performance. Cost of transport in marine iguanas
varies inversely with body mass, and foraging patterns
of various size classes appear to have been influenced
by this trend. Small marine iguanas feed on algae on
or near shore, whereas adults obtain this food by
swimming offshore and diving (Vleck et al., 1981). It
is of interest that George Bartholomew, in addition to
his other laurels, is the leading contributor to knowledge of the locomotion, thermophysiology, and metabolism of the Galapagos marine iguana, through both
the studies just cited and several others (Bartholomew
and Lasieweski, 1965; Bartholomew, 1966c; Dawson
et al., 1977; Bartholomew and Vleck, 1979).
In the latter portion of George Bartholomews professional career, he extended his research activities to
involvement in a highly productive program of investigation of insect thermophysiology. He explained this
extension thus (Bartholomew, 1982a, p. 233): . . . .
historically insect physiologists have paid relatively
little attention to the behavioral and physiological control of body temperature and its energetic and ecological consequences. Whereas many students of the comparative physiology of terrestrial vertebrates have been
virtually fixated on the topic. For the past ten years,
several of my students and I have exploited this situation by taking the standard questions and techniques
of comparative vertebrate physiology and applying
them to insects (see Heinrich, 1981). It is surprising
that this pattern of innovation is not more deliberately
employed. It is common place to find a biologist
trained in one field working in another. This represents
a more demanding change than transferring questions
and techniques between fields. The extension to in-

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tively stable population levels. The antelope ground

squirrel is a predictor, with a slow, lengthy reproductive period coinciding with the historical probability for rainfall and plant productivity, which can be
unpredictable in a given year. The Merriam kangaroo
rat (Dipodomys merriami) is a responder, breeding
in direct response to pulses in food production. Its congener, the somewhat larger Great Basin kangaroo rat
(D. microps), with a dependable food supply of salt
bush (Atriplex confertifolia) leaves, is an independent, successfully ignoring rainfall and pulses of seed
production. The lengthy periods of dormancy occurring in the two pocket mice (Perognathus longimembris and P. formosus) overlap the periods of winter
rainfall and may extend beyond it. They are in Kenagy
and Bartholomews (1985) phrase pulse gamblers
that produce large litters in a brief period, which survive in favorable times but may succumb in unfavorable years. This study further documents the existence
of multiple pathways to survival in a particular environment, even among closely related species.
Bart in collaboration with V. A. Tucker and A. K.
Lee undertook some studies on the thermal responses
of lizards which had some important results. One on
the agamid Amphibolurus barbatus provided the first
demonstration in reptiles of a physiological capacity
for modulating the rate of change of body temperature
under constant conditions (Bartholomew and Tucker,
1963), capacities that were also observed in varanid
lizards (Bartholomew and Tucker, 1964) and the large
skink Tiliqua scincoides (Bartholomew et al., 1965).
Control of heating and cooling rates was also subsequently found in the Galapagos marine iguana, Amblyrhynchus cristatus (Bartholomew and Lasiewski,
1965). In some of these lizards, though not the varanids, heart rate followed different trajectories during
cooling and heating, with the lower rates associated
with the former process and higher rates with the latter.
This probably retarded heat loss during cooling and
accelerated heat gain during warming. The role of endogenous heat production in affecting rate of change
in body temperature was considered in these studies,
but could not be resolved conclusively. Subsequently,
it was examined directly through metabolic measurements of the marine iguana during heating and cooling
(Bartholomew and Vleck, 1979). Retention of all the
endogenous heat produced by a 2.5-kg individual
basking at 308C could account for only about 5% of
the heat gain. On the other hand, with the metabolic
rate observed when the animal was cooling through
308C, complete retention would reduce the cooling rate
by 2530%. Marine iguanas tend to be more active
during cooling than basking and thus have higher rates
of heat production. Bartholomew and Vleck (1979)
concluded that these rates probably result from attempts at thermoregulatory behavior rather than a specific thermogenic response to cooling. The expertise
gained in his studies of lizards undoubtedly facilitated
Barts producing his extensive review of reptilian

GEORGE BARTHOLOMEWS CONTRIBUTIONS

flight and the production and rolling of dung balls.

Take-off and flight temperatures increased with body
mass up to about 2.5 g and were independent of mass
beyond that. These temperatures also increased with
wing loading up to about 35 N/m2, but were essentially
constant at values between 35 and 65 N/m2. The nocturnal species Scarabaeus laevistriatus often maintained a thoracic temperature of 408C or more when
ambient temperature was 258268C. The velocity of
ball rolling by this dung beetle increased linearly with
thoracic temperature from 5 cm/sec at 288C to 18 cm/
sec at 408C. For dung beetles a premium appears to
exist on rapidity of ball production and the speed with
which balls can be rolled away from areas of high
beetle concentration; this probably contributes to the
selective advantage of endothermy and the elevated
body temperatures it produces. Additionally, Scarabaeus laevistriatus resorts to endothermy during competition for elephant dung (Heinrich and Bartholomew,
1979), the resultant high body temperatures enhancing
its ability to win fights for this resource.
The range of capacities of endothermic insects is
extended still further by the detection of a homeothermic response in the elephant beetle (Megasoma elephas). It responds to ambient temperature below 208C
by increasing metabolic rate, which prevents body
temperature from falling below 208228C. The increase is not associated with any overt activity (Morgan and Bartholomew, 1982).
Work on heterothermic moths allowed comparisons
of the allometry of resting and active aerobic metabolic rates with that for reptiles, birds, and mammals
(Bartholomew and Casey, 1978). A particularly interesting fact to emerge from these comparisons was that
the scaling of oxygen consumption during flight in the
moths is virtually identical to that for bats and birds.
Detailed analyses of sphingid and saturniid moths
(Bartholomew and Epting, 1975a, b) showed that this
was also the case for mass-specific thermal conductance, though on this basis these animals were less
well insulated than the vertebrates. However, this difference disappeared when conductance of these insects
was considered on the basis of thoracic mass rather
than total body mass (Bartholomew and Epting,
1975b). A more focused metabolic comparison was
facilitated by the fact that the body masses of some of
the larger sphingid moths overlap those of hummingbirds (Bartholomew, 1981), and this provided the opportunity for a direct comparison of energetics in analogous flight systems supporting a common mode of
foraging. Both types of animals feed on nectar while
hovering, yet they have very different evolutionary
histories. An allometric analysis (Bartholomew, 1987)
based on data for hummingbirds assembled by Bartholomew and Lighton (1986b) and on those for sphingid moths available in Bartholomew and Casey (1978)
indicates that hovering costs in the two groups are very
high but virtually identical. Moreover, arthropod and
vertebrate structural and functional patterns support
similar aerodynamic efficiencies in the size decade of

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sects is not quite as simple as this narrative suggests,

for it involved some technical challenges resulting
from the miniscule size of the most diminutive insects
studied (see, for example, Bartholomew et al., 1988),
and the transient nature of several of the responses
being measured. Regarding the latter, the development
of methods for calculating instantaneous oxygen consumption of animals in open circuit metabolism systems with appreciable washout times (Bartholomew et
al., 1981), where dynamic situations such as changing
temperatures or short-term bouts of activity are involved, has been an especially useful advance.
The work carried out on insects by Bart and his
associates emphasized analysis of the endothermic capacities, thermoregulation, locomotor costs, and, in
some cases, respiration of various heterothermic species. Endothermy results primarily from intense thermogenesis in the flight muscles, with the heat largely
being sequestered in the thoracic region during warmup and activity (Heinrich and Bartholomew, 1971). It
was of interest not only from a comparative standpoint, but also because it allowed some species to be
active with high thoracic temperatures at surprisingly
low nocturnal ambient temperatures. The animals
found capable of endothermy include a large array of
moths (Bartholomew and Heinrich, 1973; Bartholomew and Epting, 1975a, b; Bartholomew and Casey,
1978; Bartholomew et al., 1981), with the smallest endothermic sphingids studied weighing only 1/15 as
much as the smallest birds and mammals (Bartholomew and Epting, 1975a); various scarab and cerambycid beetles (Bartholomew and Casey, 1977a; Bartholomew and Heinrich, 1978; Morgan and Bartholomew, 1982); several cicadas (Bartholomew and Barnhart, 1984); a tropical cockroach, Blaberus giganteus
(Bartholomew and Lighton, 1985); and the giant fly
Pantophthalmus tabaninus (Bartholomew and Lighton, 1986a). The elevated temperatures produced
through muscular thermogenesis are crucial for achieving flight in most of these insects. However, such temperatures also were observed during sustained terrestrial activity in a cerambycid and a scarab beetle (Bartholomew and Casey, 1977b), which attained rates of
oxygen consumption matching those of active mammals of comparable size. Factorial scope (ratio between rates of oxygen consumption during rest and
activity) can exceed 100 in some individuals, approximately 103 the figure characterizing homeothermic
vertebrates. The discrepancy is, of course, explained
by the fact that the latter during inactivity remain homeothermic through maintenance of relatively high
resting metabolic rates, whereas the insects become ectothermic. The factorial scopes of endothermic insects
and heterothermic birds and mammals are very similar
(Bartholomew and Casey, 1978).
In a field study of dung beetles carried out in Kenya
(Bartholomew and Heinrich, 1978), data were obtained
on representatives of several genera (in particular
Scarabaeus, Kheper, Gymnopleurus, and Heliocopris).
These animals are conspicuously endothermic during

225

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WILLIAM R. DAWSON
within a respiration chamber were determined, the two
activities involving factorial aerobic metabolic scopes
.2 and 5.7, respectively (Bucher et al., 1982). Females were found to expend more than 103 the energy
in reproduction that males do (Ryan et al., 1983),
quantifying for this frog the profound difference between the sexes in parental investment that places a
premium on female selection of high quality mates.
In another energy-oriented project, Bart participated
in an analysis of the adequacy of fat reserves, for supporting long-distance migration by fasting Swainsons,
Buteo swainsoni, and broad-winged hawks, B. platypterus (Smith et al., 1986). Based on body masses and
estimates of flight costs, basal metabolic rate, and initial fat content, it was concluded that such migration,
involving fasting over distances of several thousand
kilometers, was indeed feasible, provided that movement was achieved through soaring flight. The importance of a capacity for long-distance movements without feeding in these hawks is related to the low probability of successful foraging over unfamiliar territory
in the company of high concentrations of conspecific
individuals.
A sabbatical leave in Kenya provided Bart an opportunity to participate in a study of the energetics of
the lesser flamingo (Phoeniconaias minor), which obtains nearly all of its blue-green algal food by filter
feeding in the surface waters of alkaline lakes, principally in the Rift Valley. He and C. J. Pennycuick
(Pennycuick and Bartholomew, 1973) combined natural history data and metabolic information from the
literature to estimate the net rate at which these birds
gain chemical energy as a function of algal concentration and time spent foraging. This rate under various
conditions represents the difference between the rate
of energy acquisition through filtration of algae from
the lake water and the rate of energy expenditure due
to general metabolism and the power requirement for
the filtration. From the estimates made, a nonbreeding
lesser flamingo should be able to achieve positive energy balance if algal concentrations exceed approximately 0.12 kg/m3 of water and the bird devotes 80%
of its time to feeding. Incubation restricts foraging to
less than half the time available so an estimated algal
concentration of at least 0.25 kg/m3 of water is required. Bart and Pennycuick estimated from literature
values for cost of avian egg production that approximately a day would be needed to produce an egg at
this algal concentration. At the highest concentration
they observed, substantially less than a day would be
required to accumulate the energy for an egg. From
these considerations these authors suggested that an
opportunistic breeding strategy would be most effective: 1) where algal concentrations are high enough to
allow the flamingos to accumulate fat reserves, they
should use them to travel about investigating different
lakes; 2) when a food concentration of $0.25 kg per
m3 of water is found adjacent to a suitable breeding
site, commence breeding immediately (Pennycuick
and Bartholomew, 1973).

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110 g. Bart (Bartholomew, 1987) noted further that

the mass-specific rate of oxygen consumption in hummingbirds exceeds the highest metabolic rate recorded
in any other vertebrate. He inferred from this that both
sphingids and hummingbirds are approaching the limit
of aerodynamic performance for animals in their size
range.
Bart and associates also have conducted a number
of studies of insects that are strictly ectothermic..
Among these, geometrid moths proved of particular
interest due to the ability of some to fly at low body
temperatures (Bartholomew and Heinrich, 1973). This
capacity may be associated with the animals very
light wing loading and low wing beat frequency. Other
research on ectothermic insects involved studies of energy metabolism and locomotor costs in three species
of tropical or desert ants (Lighton et al., 1987; Bartholomew et al., 1988; Lighton and Bartholomew,
1988) and respiration and energetics of locomotion in
flightless tenebrionid beetles from the Namib Desert
(Bartholomew et al., 1985). In a field study of the
beetles (Nicolson et al., 1984), running speed of Onymacis plana averaged a remarkable 90 cm/sec (48
body lengths/sec), an apparent championship rate for
any pedestrian insect. Comparable mean running
speeds for Physadesmia globosa and Epiphysa arenicola were only 23 and 3 cm/sec, respectively. The relative speeds of the three species were found to be correlated with leg length and muscle mass, as well as
with prothoracic temperature during activity. This temperature was elevated to 36.78 and 30.58C, respectively, by behavioral thermoregulation in the diurnal O.
plana and P. globosa, whereas it probably approximated ambient temperaure (ca. 198C) in the nocturnal
E. arenicola.
Barts continuing interest in energetics has provided
a thread connecting many facets of his research. This
interest surely results from the longstanding view stated in his chapter on general energy metabolism in Gordon et al. (1982, p. 4693): The rate of energy metabolism probably integrates more aspects of animal
performance than any other single physiological parameter. Indeed, from a simplistic point of view, the
proverbial struggle for existence can profitably be
thought of as a competition for physiologically utilizable energy. We have already seen how his interest
in energetics has led, for example, to investigation of
energy conservation through various forms of dormancy in birds and mammals; to determination of the thermal dependence of metabolic scope in several lizards;
to assessment of the role of behavioral stratagems in
shaping the energy budgets of weaver finches and leafnosed bats; and measurement of locomotor costs in a
variety of insects, the marine iguana, and hummingbirds. However, he has also participated in other projects in this general area that merit mention. One of
these concerned reproductive behavior in the neotropical frog Physalaemus pustulosus. Energy costs of calling by the male and construction by a pair of the frogs
of a foam nest (including oviposition and fertilization)

GEORGE BARTHOLOMEWS CONTRIBUTIONS

tholomews research legacy. They cap a remarkable

record of scholarly accomplishment.
EDUCATION AND SERVICE
In a professorial career spanning 19471987,
George Bartholomew distinguished himself in biological instruction and is regarded at the University of
California at Los Angeles as one of the top 20 professors in the history of the institution (Anonymous,
2000). I had the privilege of being an undergraduate
student in two of his courses during his early years.
His lectures were models of clarity and he showed a
knack for linking individual facts to larger issues in
biology. The lectures dealing with what I have referred
to as experimental natural history (Dawson, 1988)
were instrumental in my decision to pursue a postgraduate career in environmentally oriented physiology. For this I shall be forever grateful.
Bart has also been a spectacularly successful mentor
of graduate students and postdoctoral scholars and has
directed the theses of 42 of the former and guided 15
of the latter. A. F. Bennett and C. Lowe (2005) have
constructed an academic geneology that includes not
only these individuals, but their students as well:
(http://128.200.122.57/login.html).
This reveals that more than 900 persons can trace their
intellectual lineage directly to George A. Bartholomew
(A. F. Bennett, personal communication), a significant
fraction of the physiological ecologists active today.
Bart has always been a very approachable, supportive,
and patient mentor, stimulating graduate students and
postdoctoral scholars to do their best through personal
interaction and his seminar courses. As is evident from
the REFERENCES section of this essay, he made a
point of involving many of these individuals in his
research program, imparting to us a concern for defining meaningful questions for investigation and for rigorously interpreting our research results. Individuals
working with him were exposed to his graceful writing
style and rigorous editorial standardsexcellent preparation for completing theses. The writing of papers
resulting from research collaborations additionally
schooled us in the publication process.
In addition to his direct interactions with students,
George Bartholomew has contributed to the educational process by producing some valuable instructional
materials. These include chapters in two textbooks,
one for introductory students in biology (Gordon et
al., 1976) and the other for seniors and even graduate
students in this field (Gordon et al., 1982). The chapters in the latter book, which is especially relevant to
the interests of integrative and comparative biologists,
deal, respectively, with the general features of energy
metabolism and with body temperature and energy
metabolism and are notable among textbook chapters
in providing valuable reference information for researchers in physiological ecology, as well as students.
Thirty-two educational films provide a further important part of the Bartholomew educational legacy. These

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I have described in the preceding paragraphs what

I regard as the major segments of George Bartholomews research on ecologically oriented physiology.
He has explored the implications of his results with
great skill and in some cases has been able to combine
his conclusions with information from the literature to
provide a firm evolutionary perspective for his work.
I shall conclude this essay with two examples of this.
The first of these dealt with the significance of bipedalism in the ecology of the protohominids. In an earlier
study of kangaroo rats (Bartholomew and Caswell,
1951), it was noted that this form of locomotion is
relatively uncommon in mammals, raising the question
of what advantage it might afford these heteromyid
rodents. They are typically associated with sparsely
vegetated habitat and must forage in the open, where
they are potentially vulnerable to a variety of predators. Bartholomew and Caswell (1951) concluded that
the special value of bipedalism to kangaroo rats arose
from the ability it imparts for rapidly changing direction in the open and thereby lowering predation risk.
Barts collaboration with the anthropologist J. B. Birdsell in an effort to reconstruct the ecology of the protohominids again raised the issue of the advantage use
of this unconventional form of locomotion would provide these ancestral humans (Bartholomew and Birdsell, 1953). The two investigators concluded that bipedalism was important to protohominids in freeing
the hands, thereby allowing continuous and efficient
manipulation of such rudimentary tools as rocks,
sticks, or bones. Bartholomew and Birdsell (1953)
therefore refined the definition of man from being a
tool-using animal (a host of other animals employ
tools) to one of being the only mammal that is continuously dependent upon tools for survival. They concluded that movement by the protohominids into this
novel dimension of behavior was importantly linked
with the advent of bipedalism.
The second example of Barts inferential abilities
resulted from the work with pinnipeds cited earlier.
This led him to wonder about factors leading to evolution of polygyny in this group, one of its most conspicuous features. He developed a model (Bartholomew, 1970) based on the special features of these animals, terrestrial partuition and offshore marine feeding, which have interacted with characteristics
common to most mammals in such away as to produce
both sexual dimorphism in size and polygynous breeding systems. Gregariousness and exclusion of most
males from females in rookeries were accorded key
roles. Also, large size and subcutaneous fat, characteristics serving to promote heat conservation during immersion of the animals, were recognized for their roles
in permitting sustained fasting and prolonged territory
occupancy by dominant males. Barts abilities to explore the full implications of the results of his research,
to maintain an evolutionary perspective, and to identify probable key steps in the evolution of particular
clades represent important features of George A. Bar-

227

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WILLIAM R. DAWSON

CONCLUDING STATEMENT
Professor George A. Bartholomew has played a major role in shaping ecologically oriented physiological
studies of animals in the period from just after World
War II until the present. His importance to physiological ecology and to the Society of Integrative and
Comparative Biology results from contributions at several levels.. An inspiring teacher, effective graduate
mentor, and innovative and productive researcher,
George A. Bartholomew has maintained a clear vision
of what can be accomplished by realizing that biology
is a continuum and has strived to work at interfaces
between disciplines. His holistic view of biology and
his recognition of the ultimate connection of organismal studies with enquiries conducted at other levels of
biological integration have made him a voice of reason
in elucidating the drawbacks for creative scholarship
of excessive disciplinary fractionation. Perhaps we can
give him no higher accolade than recognizing him as
a truly broad scholar who has had a major impact on
his many students and postdoctoral scholars as well as
on their students, on ecologically oriented physiology,
and on comparative and integrative biology generally.
The George A. Bartholomew Award provides a tangible indication of the esteem in which Bart is held by
colleagues in the Division of Comparative Biochemistry and Physiology and throughout the SICB.
REFERENCES
Anonymous. 2000. The Bruin century. UCLA Today 20(9).
Bartholomew, G. A., Jr. 1942. The fishing activities of double-crested cormorants on San Francisco Bay. Condor 44:1321.
Bartholomew, G. A., Jr. 1943a. The daily movements of cormorants
on San Francisco Bay. Condor 45:318.
Bartholomew, G. A., Jr. 1943b. Contests of double-crested cormorants for perching sites. Condor 45:186195.
Bartholomew, G. A., Jr. 1949. The effect of light intensity and day

length on reproduction in the English sparrow. Bull. Mus.

Comp. Zool. 101:433476.
Bartholomew, G. A., Jr. 1950. Reoccupation by the elephant seal of
Los Coronados Islands, Baja California, Mexico. J. Mamm. 31:
98.
Bartholomew, G. A., Jr. 1951. Spring, summer, and fall censuses of
the pinnipeds on San Nicolas Island, California. J. Mamm. 32:
1521.
Bartholomew, G. A., Jr. 1952. Reproductive and social behavior of
the northern elephant seal. U. Calif. Publ. Zool. 47:369472.
Bartholomew, G. A. 1956. Temperature regulation in the macropod
marsupial, Setonix brachyurus. Physiol. Zool. 29:2640
Bartholomew, G. A. 1958. The role of physiology in the distribution
of terrestrial vertebrates. In Zoogeography, pp. 8195. AAAS,
Washington, D.C.
Bartholomew, G. A. 1964. The roles of physiology and behaviour
in the maintenance of homeostasis in the desert environment.
In G. M. Hughes (ed.), Homeostasis and feedback mechanisms,
Symp. Soc. Exp. Biol 18:729. Cambridge University Press,
Cambridge, U. K.
Bartholomew, G. A. 1966a. Interaction of physiology and behavior
under natural conditions. In R. I. Bowman (ed.), The Galapagos, pp. 3945. University of California Press, Berkeley and
Los Angeles.
Bartholomew, G. A. 1966b. The role of behavior in the temperature
regulation of the masked booby. Condor 68:523535.
Bartholomew, G. A. 1966c. A field study of temperature relations
in the Galapagos marine iguana. Copeia 1966:241250.
Bartholomew, G. A. 1970. A model for the evolution of pinniped
polygyny. Evolution 24:546559.
Bartholomew, G. A. 1972. The water economy of seed-eating birds
that survive without drinking. In Proc. XVth Internat. Ornithol.
Congr., pp. 237254. E. J. Brill, Leiden.
Bartholomew, G. A. 1981. A matter of size: An examination of
endothermy in insects and terrestrial vertebrates. In B. Heinrich
(ed.), Insect thermoregulation, pp. 4578. John Wiley & Sons,
New York.
Bartholomew, G. A. 1982a. Scientific innovation and creativity: A
zoologists point of view. Amer. Zool. 22:227235.
Bartholomew, G. A. 1982b. Physiological control of body temperature. In C. Gans and F. H. Pough (eds.), Biology of the Reptilia,
Vol. 12, pp. 167211. Academic Press, London and New York.
Bartholomew, G. A. 1986. The role of natural history in contemporary biology. BioScience 36:324329.
Bartholomew, G. A. 1987. Interspecific comparison as a tool for
ecological physiologists. In M. E. Feder, A. F. Bennett, W. W.
Burggren, and R. B. Huey (eds.), New directions in ecological
physiology, pp. 1137. Cambridge University Press, Cambridge,
U.K.
Bartholomew, G. A. 2005. Integrative biology: An organismic biologists point of view. Integr. Comp. Biol. 45:330332.
Bartholomew, G. A. and M. C. Barnhart. 1984. Tracheal gases, respiratory gas exchange, body temperature and flight in some
tropical cicadas. J. Exp. Biol. 111:131144.
Bartholomew, G. A., A. F. Bennett, and W. R. Dawson. 1976. Swimming, diving, and lactate production of the marine iguana, Amblyrhynchus cristatus. Copeia 1976:709720.
Bartholomew, G. A., Jr. and J. B. Birdsell. 1953. Ecology and the
protohominids. Am. Anthropologist 55:481498.
Bartholomew, G. A. and T. J. Cade. 1957a. The body temperature
of the American kestrel, Falco sparverius. Wilson Bull. 69:
149154.
Bartholomew, G. A. and T. J. Cade. 1957b. Temperature regulation,
hibernation, and aestivation in the little pocket mouse, Perognathus longimembris. J. Mamm. 38:6071.
Bartholomew, G. A. and T. J. Cade. 1963. The water economy of
land birds. Auk 80:504539.
Bartholomew, G. A. and T. M. Casey. 1977a. Body temperature and
oxygen consumption during rest and activity in relation to body
size in some tropical beetles. J. Therm. Biol. 2:173176.
Bartholomew, G. A. and T. M. Casey. 1977b. Endothermy during
terrestrial activity in large beetles. Science: 195:882883.
Bartholomew, G. A. and T. M. Casey. 1978. Oxygen consumption

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analyze significant behaviors and processes of animals

and illustrate a number of biological principles. The
extensive film series on the Galapagos Islands as an
evolutionary laboratory are especially noteworthy.
Barts fairness and calm good judgment have also
allowed him to contribute important service to a number of groups. He was been Chair of the Department
of Zoology at UCLA and also participated in several
boards or committees at the national level. These include advisory panels for the National Science Foundation, the Board of Trustees of the California Academy of Sciences, and the Council of the Smithsonian
Institution. Additionally, he served as a Scientific Advisor to U.S. Marine Mammal Commission. He organized and served as Chief Scientist of R/V Alpha Helix
expeditions to New Guinea and the Galapagos Islands,
facilitating the research of groups of colleagues and
graduate students while conducting his own research.
His service to scientific societies includes terms as
Vice President of the American Ornithologists Union
and President of the predecessor of the Society of Integrative and Comparative Biology, the American Society of Zoologists.

GEORGE BARTHOLOMEWS CONTRIBUTIONS

sumption during hover-feeding in free-ranging Anna hummingbirds Calypte anna. J. Exp. Biol. 123:191200.
Bartholomew, G. A., J. R. B. Lighton, and D. H. Feener, Jr. 1988.
Energetics of trail running, load carriage, and emigration in the
column-raiding army ant Eciton hamatum. Physiol. Zool. 61:
5768.
Bartholomew, G. A., J. R. B. Lighton, and G. N. Louw. 1985. Energetics of locomotion and patterns of respiration in tenebrionid
beetles from the Namib Desert. J. Comp. Physiol. B 155:155
162.
Bartholomew, G. A. and R. E. MacMillen. 1960. The water requirements of mourning doves and their use of sea water and NaCl
solutions. Physiol. Zool. 33:171178.
Bartholomew, G. A. and R. E. MacMillen. 1961. Oxygen consumption, estivation, and hibernation in the kangaroo mouse, Microdipodops pallidus. Physiol. Zool. 34:177183.
Bartholomew, G. A. and M. Rainy. 1971. Regulation of body temperature in the rock hyrax, Heterohyrax brucei. J. Mamm. 52:
8195.
Bartholomew, G. A. and C. H. Trost. 1970. Temperature regulation
in the speckled mousebird, Colius striatus. Condor 72:141146.
Bartholomew, G. A. and V. A. Tucker. 1963. Control of changes in
body temperature, metabolism, and circulation by the agamid
lizard, Amphibolurus barbatus. Physiol. Zool. 36:199218.
Bartholomew, G. A. and V. A. Tucker. 1964. Size, body temperature,
thermal conductance, oxygen consumption, and heart rate in
Australian varanid lizards. Physiol. Zool. 37:341354.
Bartholomew, G. A., V. A. Tucker, and A. K. Lee. 1965. Oxygen
consumption, thermal conductance, and heart rate in the Australian skink Tiliqua scincoides. Copeia 1965:169173.
Bartholomew, G. A., C. M. Vleck, and T. L Bucher. 1983. Energy
metabolism and nocturnal hypothermia in two tropical passerine
frugivores, Manacus vitellinus and Pipra mentalis. Physiol.
Zool. 56:370379.
Bartholomew, G. A. and D. Vleck. 1979. The relation of oxygen
consumption to body size and to heating and cooling in the
Galapagos marine iguana, Amblyrhynchus cristatus. J. Comp.
Physiol. 132:285288.
Bartholomew, G. A., D. Vleck, and C. M. Vleck. 1981. Instantaneous measurements of oxygen consumption during pre-flight
warm-up and post-flight cooling in sphingid and saturniid
moths. J. Exp. Biol. 90:1732.
Bartholomew, G. A., F. N. White, and T. R. Howell. 1976. The
thermal significance of the nest of the sociable weaver Philetairus socius: Summer observations. Ibis 118:402410.
Bartholomew, G. A. and F. Wilkie. 1956. Body temperature in the
northern fur seal, Callorhinus ursinus. J. Mamm. 37:327337.
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