Systematics Longitudinal Studies

355

Symposium:
Longitudinal Studies of Heritability and Natural Selection
Conveners: Peter R. Grant and Fred C. Cooke

Selection During Gosling Growth on the Degree of Sexual Dimorphism

in Lesser Snow Geese
F. Cooke, D. Lank and E. Cooch
Dept. of Biological Sciences, Simon Fraser University, Bumaby, B. C. V5A LS6, Canada
Mean body size of snow geese fledging from and breeding at the La Perouse Bay colony has
declined significantly during our 24 years of study. Birds hatched at the colony at the present
time are around 15 % lighter, and have ca. 5 % shorter tarsi and culmen than those at the
beginning of the study (CoocH et al. 1991). Although there is considerable additive genetic
variation associated with body size, most of this decline could be attributed to change in an
environmental factor, namely food availability during the critical first five weeks of a gosling's
life. Gosling growth rate influenced not only size at fledging, but also asymtotic adult size.
Snow Geese show a small but distinct sexual dimorphism of body size with males about 6%
larger than females. Male and female goslings are the same size at hatch, and sexual dimorphism develops primarily due to differential pre-fledging growth trajectories. The rate of longterm decline in body size differed in the two sexes, with females declining at a slower rate than
males. This resulted in less sexually sized dimorphic Snow Geese being produced now than
at the beginning of the study. We have also found that the fledgling sex ratio has changed over
time. Sex ratios are 1 : 1 at hatch (CooK~ & HARMS~N1983). In earlier years, there was a slight
excess of males at fledging, while in recent years this has changed to a slight excess of females,
producing a significant long term change. The parallel declines in food availability, overall
growth rate, and sex ratios at fledging suggest selective norms of reaction of the two sexes are
different, dependent on the quality of the environment during the growth period. Faster growing males were favoured when food was plentiful, the slower growing females when food was
scarce. If the selective regime observed above applies within as well as between sexes, it will
impose directional selection, at this stage of the life cycle, on the genetic component of growth
rate dimorphism, with less dimorphism as an expected evolutionary response.

356

Journal fiir Ornithologie 135, 1994

Fluctuating Selection and Evolution in Darwin's Finches

Peter R. Grant and B. Rosemary Grant
Dep. of Ecology and Evolutionary Biology, Princeton Univ., NJ 08544, U. S. A.
Thirteen species of Darwin's Finches occur on the Galapagos Islands of Ecuador. They are
derived from a common ancestor which reached the islands geologically recently, possibly in
the last three million years. To understand the diversification of this group we have been studying populations on the small island of Daphne Major for the past 20 years. The aim is to
reconstruct the evolutionary history of the group from a combination of behavioral,
ecological and genetical information. The Medium Ground Finch Geospiza fort#, a
granivorous species, has been subjected to directional selection at least five times during the
past 20 years. The outstanding events occurred under contrasting conditions; in 1976-77,
associated with a drought, and in 1984--85, during the aftermath of an abundance of rain.
Measurements of six morphological traits of uniquely marked individuals, together with
repeated observations of those individuals, enabled us to determine that survival was related
to size at these times of heavy mortality. Selection gradient analysis was performed to identify
the targets of selection from among the suite of correlated morphological characters. The
drought of 1976-77 caused 85 % mortality. Large birds survived better than small ones, partly
because they were better equipped with their large beaks to crack the large and hard seeds that
were relatively (though not absolutely) common when overall seed abundance declined. In
1982-83 there occurred an extraordinary E1 Nifio event, resulting in ten times more rain than
the maximum recorded in the preceding seven years. This caused a major shift in the food
supply, towards a predominance of small and soft seeds. After extensive breeding the population size began to decline from the middle of 1984 onwards. For the next year and a half mortality (68 %) was again size selective, but this time individuals with small beaks had an advantage. Gradient analysis identified beak width, depth and length, but not body size, as targets
of selection. An association with feeding behavior is indicated by the results of a multiple
regression analysis of diet variation over a 14-year period. The proportion of small seeds in
the diet of individuals was a positive function of the abundance of those seeds and a negative
function of beak width. Natural selection occurs within generations, evolution occurs between
generations. Following each of these contrasting episodes of natural selection a micro-evolutionary change took place. This is to be expected from the known and high heritabilities of
all of the studied morphological traits. Heritabilities, determined by parent-offspring regression, are in the range 0.6 to 0.9. An evolutionary response to each selection episode was
measured by comparing the mean of each trait before selection with the mean of the offspring
of the selected parents (2). In each case the observed response was closely predicted by a formula for multivate evolution (3) which combines the effect of selection acting directly on each
trait with the indirect effect of selection on correlated traits, weighted by the appropriate
genetic variances and covariances. The correspondence between observed and predicted values
suggests that the targets of selection were correctly identified. The main implication of these
results is that the transformation of one species into another by directional natural selection
could occur relatively rapidly, given the right environmental circumstances.

Literature: 1. LANDE,R., & ARNOLD,S. J. (1983): Evolution 37: 1210-- 1226.2. GRANT,B. R.
& GRANT, E R. (1993): Proc. R. Soc. Lond. B 251: 111--117. 3. LANDE,R., (1979): Evolution
33: 402--416.

Systematics Longitudinal Studies

357

Adaptive Radiation Along "Genetic Lines of Least Resistence"

Dolph Schluter
Dept. of Zoology, Univ. of British Columbia, Vancouver, B. C. V6T 1Z4, Canada.
The effects of quantitative genetics on short-term phenotypic evolution are well supported by
theory. Studies are now beginning to use measurements of genetic parameters in natural
populations to successfully predict evolution over a single generation. The longer-term evolutionary consequences of genetic variation and covariation between traits are more controversial. This is because natural selection may determine the final outcome of evolution regardless
of genetics, unless constraints are severe (as when some traits completely lack genetic variance).
However, prior to attaining equilibrium, the path taken by an evolving population should be
biased in directions of high genetic variation, except under special circumstances. To test this
morphological differences among very young stickleback populations were measured (ca.
15.000 years old), and compared to patterns of genetic variation within one of the populations.
The two trends were closely matched: populations were differentiated mainly along the dimension that is most variable within a population (the genetic "line of least resistance"). This is
despite strong comparative evidence that natural selection is responsible for population differentiation. To test the generality of this result, similar trends in other groups were examined
from published data: sparrows, Galapagos finches, mice, and Old World flycatchers. Overall,
results show that the vast majority of evolution takes place in directions of high genetic
variability within populations. This pattern appears to be most pronounced when species are
recently diverged and weakest when more distant species are compared, although the effect of
time is noisy and needs further testing. The amount of evolution in a particular direction tends
to be less when this direction departs most from that of maximum genetic variance. This is
among the first indications that quantitative genetics can predict evolutionary trends over a
reasonably long term. This could be a reflection of genetic constraints. An alternative explanation is that the natural selection pressures that cause adaptive radiation are the same as those
which shape patterns of genetic variation within species.

358

Journal ftir Ornithologie 135, 1994

Inheritance of Phenotypically Plastic Physical Traits in the Great Tit
Andrew G. Gosler

Edward Grey Inst. of Field Ornith., Dept. of Zool., South Parks Rd., Oxford OX1 3PS, U. K.
Many physical traits of birds vary through the life of an individual, for example with season,
moult or body condition. While variation in some such traits has been shown to contain a
genetic component, a trait may present differently under different environmental conditions
so that it may be regarded as phenotypically plastic. The degree to which these 'within-individual' changes are themselves inherited has rarely been studied. Hence, for example: In the
Great Tit, although wing length has a strong genetic component, the heritability of the degree
of change in wing length with age has not been examined. This paper examines the inheritance
of seven traits which show age-related and/or seasonal change within individuals, and an eighth
which is believed to be an indicator of environmental conditions during moult, by the analysis
of data collected over twelve years in a long-term population study of the Great Tit in
Whytham Woods, Oxford. Data were collected while trapping during both breeding and nonbreeding seasons, and in many cases, parents and/or sibs were identified. Evidence is presented
that after autumn, related individuals do not associate within flocks so that, from this time,
similarities between relatives in the change in any trait are unlikely to be through a shared environment. Details of the traits concerned, and principle results are: (A) Wing length. Full
grown during the dependent period and generally shorter in juvenile than adult. The moult
responsible occurs after the first breeding season when relatives are widely dispersed on territories. Both wing length and the change in wing length across the first post-nuptial moult
were highly heritable (B) Body mass. This varies seasonally due to changes in body condition.
Although mass is heritable, seasonal changes were not. (C) Tracheal pit fat levels. This is
strongly influenced by time of day and temperature (1, 2). Neither fat levels nor seasonal or
age-specific changes in fat levels were correlated between relatives. (D) Muscle score--an index
of protein condition varying significantly through time (2). Unlike fat, this was found to be
strongly heritable. (E) Bill length. Full grown shortly after independence, this varies strongly
with season and age (3). Age-specific change in bill depth was strongly heritable but seasonal
changes were not. (F) Bill depth. Full grown shortly after independence, this varies weakly
with season and age (3). Neither age-specific nor seasonal differences were heritable. (G) Bill
index-an index of bill shape derived from bill depth/length, known to be of ecological
significance, and varying strongly with season (3). Seasonal changes in bill index were strongly
heritable, age-specific changes were not. (H) JGCs - the number of juvenile greater coverts
remaining after PJ moult, indicating conditions during moult (2). This was highly correlated
with the number of JGC retained by the father as a juvenile.
Literature: 1. GOSL~, A. G. (1987): D. Phil thesis Univ. of Oxford. 2. Gosnn~, A. G. (1991):
Bird Study 38: 1--9. 3. GOSLEt~,A. G. (1987): Ibis 129: 451--476.