Oecologia (Berl.) 37,257-272 (1978)
Oecologia
9 by Springer-Verlag 1978
Relationships between Body Size
and Some Life History Parameters
L. Blueweiss 1, H. Fox, V. Kudzma, D. Nakashima, R. Peters, and S. Sams
Department of Biology, McGill University, 1205 McGregor Avenue, Montreal,
Quebec, Canada H3A 1B1
Summary, Patterns in life history phenomena may be demonstrated by examining wide ranges of body weight. Positive relationships exist between adult
body size and the clutch size of poikilotherms, litter weight, neonate weight
life span, maturation time and, for homeotherms at least, brood or gestation
time. The complex of these factors reduces rmax in larger animals or, in
more physiological terms, rmax is set by individual growth rate. Comparison
of neonatal production with ingestion and assimilation suggests that larger
mammals put proportionately less effort into reproduction. Declining parental investment and longer development times would result if neonatal weight
is scaled allometrically to adult weight and neonatal growth rate to neonatal
weight. Body size relations represent general ecological theories and therefore
hold considerable promise in the development of predictive ecology.
A number of general, if simplistic theories have been advanced which predict
various ecological and physiological characteristics from body weight (Rensch,
1959; Stahl, 1962; Bonner, 1965; Schoener, 1968; Farlow, 1976 and others),
Such theories give average values which can serve as standards for comparison
with specifically interesting sets of data. They can form an empirical foundation
for both ecological models and for discussions of ecological generalities, and
they allow more objective evaluation of presumed trends. Such relationships
are presented as double logarithmic plots described with a power relation of
the form y = a Wb where W is body weight and a and b are constants fitted
to the data by least squares regression. In this paper, previously published
data are used to construct further theories which predict aspects of life history
and population growth from animal body weight.
Data were collected from both primary and secondary sources, but no attempt was made to survey the literature exhaustively. When available, species
1
Present address:
Dept. of Evolution and Ecology, State University of New York, Stony Brook,
New York, USA
0029-8549/78/0037/0257/$03.20
258
k Blueweiss et al.
Table 1. A comparison of equations from least squares regression analyses of the data presented in the
figures. The parameter in column 1 is described as a function of body weight (w, in grams fresh weight)
in column 4. The last column (r 2) lists the coefficient of determination which represents the proportion
of the variation in y which is explained by the regression (Steele and Torrie 1960)
Parameter
Taxonomic group
Units
Fecundity
reptiles
salmon and trout
other fish
amphibious and
aquatic poikilotherms
No./clutch
y=0.77 WT M
No. in ovariesy= 15.3 W~176
No. in ovariesy=408 WT M
No. in ovariesy=347 W~
Litter weight
mammals
birds
reptiles
poikilotherms
g/litter
g/clutch
g/clutch
mma/clutch
y=0.55 14/o.82
y = 1.24 W0"74"
y=0.35 WT M
mammals
birds
birds
reptiles
fish
Crustacea
fish and Crustacea
g/neonate
g/egg
g/hatchling
g/egg
mm3/egg
mm3/egg
mm3/egg
y=0.097 14/0.92
y=0.26
W 0"77
y=0.28
y=0.41
y=0.59
y=0.01
W0'69
W~
WT M
W~
y=0.06
W 0"77
0.94
0.83
0.86
0.70
0.26
0.35
0.82
rmax
virus to mammals
day-1
y=0.025
W -~
0.90
Defecation
mammals
g/g/day
y=0.85 W -~
Neonate production
mammals
g/g maternal y=0.037 W 0.43
body wt/day
0.65
Brood time
birds
days
y=9.1 W~
0.47
Gestation time
mammaIs
days
y = 11 W~
0.72
Average life span
mammals
days
y = 630 W~ 1v
0.56
Maturation time
virus to mammals
days
y=53 W~
0.96
Neonate weight
Equation
r2
y = 2 7 5 WT M
0.78
0.69
0.78
0.58
0.97
0.85
0.96
0.86
0.80
means were recorded, but if necessary the midpoints of ranges were used. Data
were fitted to a power relationship using a pre-programmed analysis provided
with a Hewlett-Packard 97 desk-top calculator. This program provided intercept
(a), slope (b) and the coefficient of determination, r 2 (Steele and Torrie, 1960).
Such an approach has several short comings. Because the surveys are not
exhaustive, particular groups may exert an undue influence on the relationship.
Reliance on secondary sources could introduce biases, both conscious and unconscious, of other scientists. Regression analyses are strictly valid only if the
variance in Yis independent of X and is normally distributed about the regression
line, but this analysis is usually applied even if these requirements are not
met. In keeping with this practice, ordinary predictive regressions in which
b = S x y / Z x 2, where x = X - X" and y = Y - Y have been applied here. Ricker
(1973) suggests that a functional regression in which slope is (Sy2/Zx2) 1/z
is preferable for body weight relations, but this statistic has fewer adherents.
The slope of the functional regression for the curves presented here can be
calculated as b/r (Ricker 1973) using the values listed in Table 1. This increases
Body Size and Life History Parameters
259
IO
$4"
LITTER
It
WEIGHT
in
,r e
"'*
** s
MAMMALS
I00
I0
..~ .i'...b-"...~
I k(J
...:.
9
lO0
"-~":;: ""
y = o.as . 8 2
9
lO
I
Ig
I0
IO0
Ikg
I0
BODY
I00
It
IO
I00
WEIGHT
Fig. 1. Mammalian litter weight (y, in g per litter) vs maternal body weight (W, in g). Data
from Sacher and Staffeldt (1974) and Leitch et, al. (1959)
/
I0 4
#
/
/
9 ~
10 3
#
~ 1 4 9##
9
".";',~.'~'.F
9 .:.'*,~r~'~'"
-.r
;,
~io2
o
..
.&kq,.=,. 9
l-"r
"~-~,
,
Zo,,
~
c=,..w-
9
b-
9 "~,'~
.J
q~
~149
##" 9
9
~
t
" ~
~
/
BIRDS
data from HEINROTH
i
Io
i
ioz
BODY
i
i
i
Io3
Io4
io~
WEIGHT (W) grams
Fig. 2. Total weight of avian clutch vs maternal body weight
260
L. Blueweisset al.
i0 ~
103
iO 2
9. :: .;.
::!:...------..
LU
m cc
o
> lOB
I
cD
I0
J
o
I0
,c~
~
Y = 275
W 's2
OI
rnrn 3
I
I mg
I0 mg
I
IO0 mg
I
I
I
I
Ig
lOg
IOOg
I kg
BODY
I
I0 kg
I
IOOkg
WEIGHT
Fig. 3. Clutch volume of poikilotherms (frogs, reptiles, fishes and crustaceans) (y, in mm3) as
a function of maternal body weight (W, in g). A specific gravity of 1 was assumed in order
to convert weight to volume for frogs and reptiles. Sources as in Figures 7 and 8 plus Skrzypiec
(1964)
the slope but has a pronounced effect only when the scatter in the data is
large. In any case, equations with low r 2 values should be viewed with caution.
The first Figures (1-3) show that litter weight is tightly correlated with
maternal weight (Sacher and Staffeldt, 1974; Millar, 1977) for all groups examined. G o o d relations also exist between the weight of individual offspring and
maternal weight (Figs. 4-8). Poikilotherms increase fecundity per clutch as adult
body size increases (Figs. 9 and 10). Certain anecdotal information (Southwood
et al., 1974) suggests that this may also hold for homeotherms, but neither
birds (data from Heinroth, 1922) nor mammals (data from Leitch et al., 1959;
Sacher and Staffeldt, 1974) showed a relationship between the number of
offspring per litter and adult body weight. Millar (1977) previously observed
this lack of relation in small mammals. The average clutch size of birds in
Heinroth's (1922) data is 4.85 (standard deviation=3.04) and, for mammals,
the mean number per litter (Leitch et al., 1959; Sacher and Staffeldt, 1974)
was 2.71 (S.D. =2.19).
Comparison among these regression lines (Fig. 11) shows that the litter
weight of most animals is a strikingly similar function of adult body weight.
Homeotherms of similar adult body size also produce offspring of similar size,
but individual egg size for reptiles increases less rapidly with adult body size
reflecting the poikilothermic tendency to increased clutch size in larger animals.
Fish and Crustacea clearly produce more young per unit of litter weight, even
though the scatter in Figure 9 is quite high. Figure 11 shows that smaller organ-
Body Size and Life History Parameters
261
/
,hI05
~
r
/
,~
9 o
x
104
<.o
~103
0
.j
<lo
2
Y = 0 . 0 9 7 W0 9 2
g
>_-Io
_z
~
*~
~
MAMMALS
Lc
SACHER +STAFFELDT
9 LEITCH~HYTTEN+BILLEWICZ
S9
O
,,, o j
S9149
i
i
lgram I0
i
=
3
'4
5
I0
10
IO
BODY
WEIGHT
I0
i
1ton
(W)
!0
tO0
Fig. 4. Weight of individual neonate (y, in g) vs maternal body weight (w, in g) for mammals
I000
AVIAN
NEONATE
9. 9
WEIGHT
sS 9
sS
ioo
ss
9SsSI
9
,1o1~
I ~
,-,-..
=
9
9
,
! ! I!
...:.;:...-
Io
t I I
i'-
I I
W
s~
r
z
#o
I
#0
J
,"~':"
Y =0.28
W "69
so
s~s Sr
Ig
10
MATERNAL
I00
BODY
I kg
10
WEIGHT
Fig. 5. Individual hatchling weight (y, in g) vs maternal body weight (w, in g) for birds. Data
from Heinroth (1922)
262
L. Blueweiss et al.
/
/
/
/
IO~
9
/
;'.."
.
/.
, /" '
E = .Z6 w .77
i0 ~
9
I0
I"r
9 ~. :L
;.,}}.,.
.~''
B,Ros
bJ
dclt~a from HETNROTH
/.
i
I0
n
i
102
BODY
a
,
I0 S
104
105
W E I G H T (W) g r a m s
Fig. 6. Egg weight (E, in g) vs maternal body weight for birds
EGG
r~
E
E
SIZE
fogether
I000
(D
o
uJ
w 77
sS
I00
..J
o
>
y=O.06
9
9
[O
o9
:..
9149
*e
9
r
"~,,'/
rJsh
9 ."~g" ~ " ' "
y=0.59
See
w
I
/
j.o,, 9 9 9 . .
-
O.I
es
9
~e
s ~
>
O.OI
Z
0.001
~
ling
o
o ,,,,~ustaceans
~ - ~
z
I0
~o
o
I00
~A
y =0.01 w ~'~
I g
BODY
I0
I00
I kg
I0
100
WEIGHT
Fig. 7. Individual egg volume (y, in mm 3) vs maternal body weight (w, in g) for fish, crustaceans
and for both. Data from Scott and Crossman (1973), Carlander (1950), Colette and Earle (1922),
Juszcyx (1971), Kozlowska (1971). Abrahamsson and Goldman (1970), Apollonio (1969), Mauchline
and Fisher (1969)
Body Size and Life History Parameters
REPTILIAN
Ioo
EGG
y = 0.41
263
WEIGHT
w 0"42
(.9
(.9
uJ
I
I0
I00
IO00
MATERNAL
BODY
I0000
WEIGHT
(g)
Fig. 8. Individual egg weight (y, in g) vs maternal body weight (w, in g) for reptiles. Data from
Bustard (1971), Goldberg (I971), Parker (1973), Vitt and Ohmart (1974), Parker and Pianka (1973),
Tanner and Krogh (1974), Vitt (1975), Clark (1974), Limpus (1973), Graham (1971), White and
Murphy (1973), Yntema (1970), Cagle (1950, 1952)
CLUTCH
SIZE
OF
AQUATIC
POIKILOTHERMS
o
o
,.,
.J
<
I0 6
,.
LL
(3C
,,,
o_
10
o
5
o
o
o
frogs
o
i0 4
y=347
c~c~
(.9
LIJ
d
z
fish
crustaceans
w
0.47
I03
o
o
9 1 I9
9
"~"
9-" o 9
,,~,
o
o
o
o ~,
o~176 o
.o o o
a
o
%
~
o
o
~149
o
8
o~% .,.,o'~176176
%o 0oo%~176
o
o
o
oooo
o~
oo
o
o
o
Oo
.,o
o
10 2
8
o
I0
I00
Irng
IO
BODY
I00
Ig
I0
I00
I kg
I0
I00
WEIGHT
Fig. 9. The influence of maternal body weight (W, in g) on the number of eggs in the ovaries
or in the clutches of aquatic and amphibious poiklilotherms. Sources as in Figure 7 plus Skrzypiec
(1964)
isms invest m o r e in r e p r o d u c t i o n , relative to their b o d y weight, a n d that, for
a n i m a l s o f a given size, this i n v e s t m e n t is r o u g h l y a c o n s t a n t f r a c t i o n o f their
weight. Conversely, n e o n a t e s o f smaller a n i m a l s are closer to the a d u l t b o d y
size; the difference between a d u l t a n d n e o n a t e size is m o r e p r o n o u n c e d in
p o i k i l o t h e r m s , especially large p o i k i l o t h e r m s . A m o n g m a m m a l s , the ratio o f
264
L. Blueweiss et al.
REPTILES
y : 0.77
I000
w
0.48
o ' J
I00
Ig
I0
I00
Ikg
BODY
IO
I00
WEIGHT
Fig. 10. The influence of maternal body weight (W, in g) on the number of eggs per clutch in
reptiles. Sources as in Figure 8
t//
o
LU
/
Ig
_.1
p0
I--
/
Zlkg
Ikg
--BIRDS
MAMMALS
-----..........
- - - - -
/
DA
/
/
/.
~
d
/
/
R'E P T I L E S
FISH 8=
CRUSTACEA
Ig
a
>
Z
Ig
Ikg
It
MATERNAL
Ig
BODY
lkg
It
WEIGHT
Fig. 11. Comparison of regression lines of clutch weight vs maternal body weight and of egg
or neonate size vs maternal body weight for birds, mammals, reptiles and fish and Crustacea
neonate weight to maternal weight changes less rapidly with changes in maternal
weight.
Rates of reproduction are also related to body size, although less variance
is explained in these cases (Table 1). Figure 12 summarizes the relationship
between mammalian gestation time and body size; Fig. 13 gives data regarding
the length of brood time in birds. While it is hardly surprising that the larger
neonates of larger animals require longer developmental times, the exponents
in these relationships suggest that the amount of resources available to the
neonate is set by adult body size and that the rate of utilization of these resources
is determined by neonate size (see below).
Body Size and Life History Parameters
I000
MAMMALIAN
265
GESTATION
~s0S
TIME
u)
s"
ee #el.O#'
o
9 e%SS
: " a ?
v
9
la.J
sS
.
9
tl#'.
9
esS
,,* 9
"
~.s
9 9
.K
t
9
?, 9
.s"
z
0
I-,<
I-r
W
e9 9
9
I00
l.-
9 ~
y=ll
w 0'26
!
(Saeher
&
Staffeldt)
9
@#~i, 9 1 4 9 1 4 9 9
,r S
9
I0
Ig
I0
I00
I kg
I0
BODY
Fig. 12. M a m m a l i a n
I00
It
IO
I00
WEIGHT
g e s t a t i o n t i m e (y, in d a y s ) as a f u n c t i o n o f m a t e r n a l b o d y w e i g h t (w, in
g)
~100
:
:
9
I0
9
"f,.."i"
,
".:. I
g
: .. 2:;:.: . . . . . .
.~:~:,2~,~i%'~,
9
'" "
T
" ..........
~ .....
9.11WAG
BIRDS data from HEINROTH
o
BODY
WEIGHT
W
groins
Fig. 13. Avian brood time (y, in days) as a function of maternal body weight (w, in g)
G r o w t h and development are also functions of adult body size. Over a
very wide range of organisms (i.e. from virus to whales), the time to sexual
maturity is a function of body size (Fig. 14). Bonner (1965) took this parameter
to be an estimate of generation time as well. The m a x i m u m life span also
increases with body size (Rensch, 1959) in a miscellaneous group of animals
(Daphnia
to elephants, Fig. 15), as does the average life span in m a m m a l s
(Fig. 16). Since the inverse of life span approximates the death rate in the
absence of predation and disease, these figures may show that physiological
death rate increases as body size decreases.
266
L. Blueweiss et al.
TIME
TO
MATURITY
IOO,OOO
>-
IO,OO0
E3
..t
. 9
O.o~
I000
.'.'...-e
.
Ld
9 7r
~I-
ioo
z
o
I0
"~
1,0
D
0 ,[
F-
,01
,r
.
/
o~
g.."
"P' "
~,
Y= 5 5
W'
,/
2//'"
I
I
Ifg
I
I
I
I
Ipglng luglmglg
I
I
Ikg Iron
WEIGHT
Fig. 14. Maturation time (y, in days) as a function of adult body weight (w, in g) for a broad
range of animals. Data from Yarwood (1956), Altman and Dittmer (1962), Bonner (1965), Chapman
(1928), De witt (1954), Reay (1970), Demir (165), McCullough and Inglis (1961), Belynina (1969),
Burton (1962), Raitt (1968), Meuller et al. (1976), Kirkpatrick (1944), Blacker (1971), Keith (1960),
Johnsgard (1973), Weller (1965), Elder (1946), Schreiber (1976), Rue (1969), Stonehouse (1975),
Williams (1963), Rutter (1962), Rutter and Pimlott (1968), Perry (1970), Cowie (1966), Warmer
(1966), Caldwell and Caldwell (1972) and Batchelor (1963)
MAXIMUM
LIFE
SPAN
9 MAMMALS, INVERTEBRATES,
REPTILES,AMPHIBIA
+ BIRDS
,,I, FISH
Z
,,~
Y = 1570
I00,000
W "15
13_
co
lO,O00
LIJ
co
>~
I000
lOO
D ~
z;
x
IO
I
I
Oolmg ling
I
I
I
IOmg IOOmg Ig
BODY
I
lOg
I
I
IOOg Ikg
I
I
1
IOkg IOOkg II"on
WEIGHT
Fig. 15. Maximum life span (y, in days) for a broad range of animals vs adult body weight (w,
in g). Data from Grzimek (1974), Walker (1975), and Altman and Dittmer (1962, 1973)
Body Size and Life History Parameters
267
cO
AVERAGE MAMMALIAN
LIFE SPAN
v
Y = 650
5
or)
I0,000
i.d
LL
w
17
.. ,)'..:".'.&-.- r
I000
,,,,,,e.---l"~
9
~ 9
9
"*
I O0
1 ~g
I mg
BODY
I :k g
WEIGHT
I
~on
Fig. 16. Average life span (y, in days) of mammals vs adult body weight sources as in Figure 17
I 000
I00
.,,,.,,,..
",,,~
",,.
[0
OJ
INTRINSIC
.
'~"'".'. ,,.
"
Y
= 0.025
RATE
0 F
"*"~
W
NATURAL
-,26
~'~'~
"~r
INCREASE
9
O.Ol
"+
9
0.001
" *.~149
I fg
I pg
I ncj
BODY
I pg
I mg
Ig
I kg
I ton
WEIGHT
Fig. 17. The intrinsic rate of population growth, r~,, (y, in day t) vs adult body size (w, in
g) for a broad range of organisms. Data from Fenchel (1974), Nagel and Pimentel (1964), Clarke
and Sardesi (1959), David and Fouillet (1971), Frazer (1972), Wrensch and Young (1974), Lauglin
(1965), Murphy (1967), French and Kaaz (1968), Watson (1970) and Lowe (1969)
Smith (1954) and Fenchel (1974) have shown that the intrinsic rate of population growth, r~,a,, decreases with adult body size. Fenchel (1974) interpreted
his data as three parallel curves corresponding to the rm,x of unicells, poikilotherms and homeotherms. However, reanalysis of his data showed that all
points could be treated with a single curve (Fig. 17). The product of rmax and
adult weight is an estimate of the maximum possible rate of production. Figure 18 compares this transformation with the observed rates of individual growth
reported in a range of ecological studies (Ecology 1961 to 1975, J. Anita. Ecol.
1968 to 1975; Peters, 1978). The close similarity between the curve and the
data points suggests that differences between rm~ and observed rates of population growth may reflect increased predation rates rather than decreased rates
268
L. Blueweiss et al.
!
HOMEOTHERMS 9
POIKILOTHERMS o
. ~ 103
1
-6 {o 2
E
o
I0 i
9
..1
1.
E
o
t.
lO_i
,r
rr
"II.-
IG 2
o
~IG3
IT"
~176
~176
o
o
o/
~
~176176
o
0.74
_
rm= x =0.025
o ~ Wx
W 0"7
~ o ~o
iO-4.
o
10"5
i0-4
i0-3
BODY
I0 -2
i0-I
WEIGHT
i01
(W) grams
102
i0 :~
Fig. 18. A comparison of m a x i m u m rate of population growth in grams individual- 1 day 1 calculated as rma, • adult body weight (curve and equation) and measurements of individual growth
rates as reported in Ecology 1961 to 1975 a n d J. Anim. Ecol. 1968 to 1975 (data points for
h o m e o t h e r m s and poikilotherms)
I000
MATERNAL
IN
I00
g
LO
~E
O.I
PRODUCTION
MAMMALS
vs
AVAILABLE
I0
>-
NEONATE
ENERGY
I = INGESTION
RATE
D= DEFECATION
R=RE;:::T,O"
IRATE
(.9
0.01
<D
v
LU
<I
O, O O i
0.0001
0,0000(
=NE NA E
IG [//MATERNAL WEIGHT//'GESTATION TIME
y
0.000001
I
.01
= 0,057
I
O,I
L
1.0
BODY
W -'45
t
IO
I
I00
I
I000
I
I0,O00
WEIGHT (KILOGRAMS)
Fig. 19. C o m p a r i s o n of energy available from ingestion or assimilation and the rate of production
of neonates (y, g of neonate/g of maternal weight/day) over a range of maternal weights (w,
in g). Data from Sacher and Staffeldt (1974)
Body Size and Life History Parameters
269
of birth and individual growth. However, much of the data regarding individual
growth rates was obtained in the laboratory and might over-estimate natural
rates.
For many organisms, metabolic rate per animal varies as W~ (Hemmingsen, 1960). This implies that metabolic rate per gram varies as W -~
and
that the time required for a unit of metabolic work varies as W ~
(Stahl
1962). Table 1 shows that exponents from equations describing growth and
reproduction are frequently similar. Some of the departures from these exponents
probably reflect only scatter in the data, but others may be real.
Among the most interesting of the apparent departures is neonatal production, defined as litter weight/(maternal weight x gestation time). Figure 19 compares these data with the energy available per gram from ingestion (from Farlow,
1976) or assimilation defined as ingestion-defecation and as ingestion-(defecat i o n + b a s a l metabolic rate) (Hemmingsen, 1960). Apparently larger mammals
put less effort into reproduction than do small relative to available energy.
Millar (1977) reached a similar conclusion based on the metabolic demands
of the litter just prior to weaning. His calculations show that the effort of
the mother declines as W o.,.2, a value very close to that in Figure 19, W - ~
Pearson (1968) observed that small sea birds spend a larger proportion of
their time finding food for their young than do larger sea birds; this may
be a specific example of the increased cost of reproduction in smaller homeotherms.
The exponent, - 0 . 4 3 , can be rationalized if the rate of neonate production
is considered a function of neonate body weight, W,, rather than maternal
body weight, W. Since
W, =0.097 W ~
(1)
and production, Pn in g/g of neonate/day is approximated (Fig. 17) by
P, =0.025 W2 0.26
(2)
the average rate of production of neonates, P in gig of maternal weight/day,
can be determined by substituting Equation 1 for W, in Equation 2 to give
P, as a function of maternal weight, and then multiplying by (total litter weight/
maternal weight)
P=PnxO.55 W~
W 0.,,2
(3)
A parallel argument can be applied to gestation time (G) and brood time
(B). Since G is the time required to produce a neonate, then G can be calculated
as neonate weight divided by foetal growth rate; substituting Equation 1 for
W, gives G as a function of maternal weight, W
G = W,/0.025 W ~
W~
(4)
and B can be predicted from hatchling weight, WH
B = W~/0.025 17/O"74=29 W 0-18
(5)
270
L. Blueweiss et al.
The exponents predicted by assuming that foetal growth rate is a function
of neonate size and that neonate size is a function of maternal weight are
quite close to those observed when the scatter in the data and the approximation
involved in substituting neonate weight for some average foetal weight are
considered. The elevations in Equations 3 to 5 are higher than those observed
(Table 1) which suggest that actual embryonic growth rates are two to three
times faster than those predicted from population growth rates (Eq. 2). It is
a commonplace that individual growth rate decreases with age (Brody, 1945;
Millar, 1977) if the rate is expressed per unit weight.
Each of the curves presented in this overview of the life history and reproductive processes is a general biological theory (Wilkie, 1977). Each predicts some
general characteristic which is difficult to measure, from a simple, easily measured, independent variable, body weight (Peters, 1977). By comparing curves,
we can identify ecological constants like total investment/clutch (Fig. 11) and
regular shifts such as the demands of reproduction relative to available energy
(Fig. 19). The most likely applications of such theories are in treatments of
community production (Sheldon et al. 1977), succession, material flow (Peters,
1978) or other properties of multi-specific animal assemblages. However even
a confirmed autecologist will find these curves more suitable standards of comparison than data from individual species. Lastly, these relationships illustrate
both the strengths and weaknesses of biological trends which many of us presume
but frequently cannot adequately demonstrate.
Acknowledgements. We wish to thank Ms. P.A. Horton for her assistance in the preparation of
this manuscript.
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Received April 10, 1978