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REDWING
BLACKBIRD
YELLOW
WARBLER
GARTER SNAKE
BRONZE SONG
GRACKLE SPARROW
SNAIL
160
by Robert M. May
S
trictly speaking. ecological systems ber of species. more detailed questions vidual mangrove trees. Drawing on that
as such do not evolve. As has been can be asked about the patterns of com evidence. P. J. Darlington's Zoogeogra
stressed repeatedly in this issue. munity organization. What accounts for phy: The Geographical Distribution 0/Ani
natural selection acts almost invariably the relative abundances of individuals mals. a classic study of the past and pres
on individuals or on groups of relat among the various species in a given ent whereabouts of the vertebrates. con
ed individuals. Populations. much less region? Why do some communities cludes: 'Throughout the recorded histo
communities of interacting populations. include several very common species ry of vertebrates. whenever the record is
cannot be regarded as units subject to along with a few rare ones whereas in good enough. the world as a whole and
Darwinian evolution. It nonetheless re other communities the individuals are each main part of it has been inhabited
mains true that the forces shaping natu distributed in roughly equal numbers by a vertebrate fauna which has been
ral selection among individuals involve among the species? Why do virtually all reasonably constant in size and adaptive
all manner of biological interactions communities have many more species structure. Neither the world nor any
with other species: flowering plants with of little animals than of big ones? Why main part of it has been overfull of ani
their pollinators. fruits with the animals do food chains typically have only three mals in one epoch and empty in the next.
that disperse their seeds. distasteful in or four levels (plant. herbivore. first car and no great ecological roles have been
sects with the species that mimic their nivore second carnivore). in spite of the long unfilled. There have always been
warning coloration. Therefore in a sense fact that there is great variability in the (except perhaps for very short peri
constellations of species can be viewed amount of energy flow and in the physi ods of time) herbivores and carnivores.
as evolving together within a conven cal details of the organisms involved? large and small forms. and a variety of
tional Darwinian framework. There is The answers to such questions are different minor adaptations. all in rea
nothing of the crude notion of "group not only interesting in themselves but sonable proportion to each other. Exist
selection" in the recognition that evolu also important for conservation and re ing faunas show the same balance. Ev
tion produces patterns at the level of source management. Notwithstanding a ery continent has a fauna reasonably
ecological systems. Such patterns are growing accumulation and synthesis of proportionate to its area and climate.
anchored in the interplay of biological empirical evidence. however. there is at and each main fauna has a reasonable
relations that act to confer specific ad present no consensus on the answer to proportion of herbivores. carnivores.
vantages or disadvantages on individual any of the above questions. I shall de etc. This cannot be due to chance."
organisms. a concept that has been cap scribe here a number of the empirical Similar evidence for a variety of fossil
tured memorably in the title of a book patterns that have emerged and review groups is presented elsewhere in this is
by G. Evelyn Hutchinson: The Ecologi some speculations that have been put sue [see "The Evolution of Multicellular
cal Theater and the Evolutionary Play. forward about the underlying causes. Plants and Animals." by James W. Val
How do evolutionary forces combine entine. page 140]. One particularly nice
with the physical environment to shape vidence for consistent patterns in the example is provided by a comparison of
a community of living things? Why. for E number of species associated with a the number of families of land mam
example. does Britain have about 60 given region comes from sources rang mals in North America and South
resident species of butterflies whereas ing from the grand sweep of the fossil America before. during and after the
New Guinea has close to 1.000? Going record to controlled experiments on the formation of the Panama land bridge
beyond the question of the total num- arthropod fauna associated with indi- between the two continents in the Pleis
tocene epoch some two million years
ago. Prior to the formation of this link
the two continents had had no direct
contact since the dawn of the age of
IDEALIZED FOOD WEB, shown in the drawing on the opposite page, maps out what eats mammals. and even before that the con
what in a willow forest in Canada. Such food webs can be roughly organized into a hierarchy
nection was circuitous. by way of Eu
consisting of a small number of "trophic levels" (in this case four), although usually not all the
rope and Africa. Initially no families of
species present fit neatly into the classification scheme. Here the trophic levels are identified
land mammals were common to both
as primary producers (the willows themselves), herbivores (a variety of insects), first carnivores
(spiders and frogs) and second carnivores (birds and snakes), The pathway from plants to
continents. and given ecological roles
snakes, however, can be traced either through two intermediate links (insects, frogs) or through were played by phylogenetically distinct
three intermediate links (insects, spiders, frogs); similarly, the patbways from plants to birds actors in North America and South
can go either directly through one link (insects) or indirectly through two (insects, spiders). America [see illustratioll Oil next page].
161
0'
8'
A FULL CONTINENTAL COMPLEMENT of land mammals corresponding South American marsupial carnivore (8'), the North
evolved independently in both North America and South America American camel (C) and the South American camel-like Iitoptern (C),
prior to the formation of the Panama land bridge during the Pleisto the North American horse (D) and the South American horselike li
cene epoch some two million years ago. As a result given ecological toptern (D'), the North American rhinoceros (E) and the South Amer
roles were played by phylogeneticaUy distinct actors on each conti ican toxodont (E'), the North American chalicothere (F) and the
nent. For example, a number of such evolutionarily convergent types South American homalodothere (F'), and the North American saher
of mammals are identified by the lettered pairs. They include the toothed cat (G) and its South American marsupial counterpart (G').
shrew family in North America (A) and the caenolestine marsupial After the present land connection was formed sometime in mid-Pleis
family in South America (A'), the North American wolf (B) and the tocene, a great faunal exchange ensued between the two continents.
162
163
i
en i
whether or not a particular population is
likely to persist. At one extreme there
w en
w are species whose mortality patterns are
!;;: !;;: determined almost entirely by the vicis
a:
a:
z situdes of an environment that to them
o z
o is unpredictable, transient or patchy.
a: f=
u For them the evolutionary pressures are
(!) z
f= to produce many young, to invest little
x
w in parental care (since it has scant effect
on survival probability) and to spread
the risks by high dispersal ability. Such
"boom and bust" species are the pio
neers in the world of plants and animals.
At the other extreme are species whose
mortality is predominantly influenced
by interactions with their own species
and other species. For them the evolu
tionary pressures are to be a good com
NUMBER OF SPECIES PRESENT petitor and to have few offspring but to
invest more time and energy in raising
DYNAMIC-EQUILIBRIUM MODEL of island biogeograpby, formulated by the late Robert them. These are of course the two ex
H. MacArthur and Edward O. Wilson, holds that in both real and virtual islands local extinc tremes of a continuum. and most species
tions will on the average be compensated for by fresh immigration of the same species or other
occupy some intermediate position.
species. Extinction rates (colored curves) will tend to rise as the total number of species on the
Practical consequences attend this
island rises, and this effect will be more pronounced on small islands than on big ones. Converse
recognition of the variety of possible
ly, the rate of immigration of new.species (black cun'es) will tend to fall as the number of spe
cies rises, with the immigration rate being lower for islands far from the main species pool than life-styles. For example, as has been
for nearby ones. It follows that the equilibrium number of species present in a given area will stressed by T. R. E. Southwood of the
be highest on the large, close islands (poill1 A) and lowest on small, distant islands (poillt B). Imperial College of Science and Tech-
164
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SUMMER
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165
125
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153
155
MOUNTAINS OF NEW GUINEA (color) represent a kind of island number of highland bird species counted on each small mountain
archipelago for the species of birds that are confined to higher eleva range and at three different locations on the central range. In keep
tions. For such species New Guinea in effect consists of a large cen ing with the dynamic-equilibrium model of island biogeography the
tral island habitat (or mountain range ) and about six smaller island larger virtual islands generally have more species. This area effect
habitats along the northern coast, separated from one another by a explains most of the variation in the observed number of species;
"sea" of intervening lowlands. The numbers on the map indicate the the residual variation is largely correlated with variation in altitude.
166
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167
1 68
BUSHYTAIL
WOOD RAT
UINTA
CHIPMUNK
GOLDEN-
MANTLED
GROUND
SQUIRREL
PRAIRIE
VOLE
YELLOW-
BELLY
MARMOT
NORTHERN
POCKET
GOPHER
VAGRANT
SHREW
NORTHERN
WATER
SHREW
WESTERN
JUMPING 33
MOUSE
PIKA 121
SHORTTAIL
WEASEL 58
BELDING
GROUND 382
SQUIRREL
WHITETAIL
2,500
JACKRABBIT
PATTERNS OF EXTINCTION of 13 small mammalian species re "nonequilibrium insular biogeography," since ''the mammalian fau
sult from the relaxation of the supersaturated faunas of the 17 Great nas of the mountaintops are true relicts and do not represent equilib
Basin mountain habitats toward the smaller species numbers appro ria between rates of colonization and extinction." The colored cells
priate to particular mountaintops. The mountain ranges are named denote the locations of the surviving species. (The 13 species shown
at the top, along with the numbers that indicate their location on the in the illustration are not depicted to the same scale; their relative
map on the opposite page. Brown refers to this case as an example of sizes are indicated by giving a typical body weight for each species.)
169
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SPECIES IN ORDER OF ABUNDANCE
. 01 L-------------------------------
SPECIES IN ORDER OF ABUNDANCE
PATTERNS OF RELATIVE ABUNDANCE of various species in few species are dominant to one in which there is a large "middle
two different kinds of ecological community are contrasted. The se class" distribution of individuals among many species. The series of
ries of graphs at the left is typical of an early successional communi graphs at the right is typical of a mature ecosystem that has been se
ty, in this case an abandoned agricultural field in southern Illinois verely disturbed in this case an experimental grass plot at the Roth
that has been studied by Fakri A. Bazzaz of the University of Illinois. amsted Experimental Station in England to which nitrogen fertilizer
The plant species present in the field were counted at five stages, has been continuously applied since 1856. An analysis conducted by
ranging from one year to 40 years after the fields were abandoned. W. E. Brenchley, K. Warington, R. A. Kempton and R. L. Taylor of
The observed relative-abundance patterns are expressed in terms of the Rothamsted Station shows that the resulting relative-abundance
the percentage each species contributes to the total area covered by patterns, measured at sporadic intervals over the following century,
all the species in the community, plotted against the rank of the spe look like a normal early-successional sequence that has been run back
cies, ordered from the most abundant species (left) to the least abun ward. Presumably the disturbance had the effect of distorting the or
dant (right). The gray bars are herbs; the white bars are shrubs; the ganization of this pasture community by emphasizing one ecological
colored bars are trees. As in most such communities the overall trend factor at the expense of all the others, leading to the strong domi
is from dominance to diversity, that is, from a situation in which a nance of a single grass out of the three surviving species of grasses.
170
GreatWine.
mentary state. As Ernst Mayr has put
it: "We must take it for granted that a
large part of the mite fauna of the world
will remain unsampled. unnamed and
unclassified for decades to come." This
Or GreatWhisky.
could easily mean that the classes of ani
mals smaller than a centimeter in length
are underestimated in the illustration by
a factor of two or more. Another diffi
culty. which might account for the de
crease in numbers of species of very
small size. is that conventional taxo
nomic concepts have dubious validity
once one goes below the one-millimeter
size class.
Setting aside these caveats. the overall You can prove Old
trend for organisms ranging in size from Forester has a unique
about a centimeter to a few meters is
that a threefold increase in length corre
taste without even
sponds roughly to a tenfold decrease in putting it to your lips.
the number of species. Part of the expla
nation of this trend is clearly that small
T he same way experts
animals can subdivide the habitat more prove it with wine,
than large ones can; a species of small
plant may be merely one among many
by color.
in the diet of a large herbivore. and yet it For example, when
may provide a rich variety of different
niches wherein several species of small
held to a candle, a Burgundy should glow a deep
arthropods can coexist. purple. A Bordeaux, brick red.
Two different colors, two different tastes.
ithin a given community the bio
W logical relations among species When held to a candle, a great whisky should
can be depicted as a food web in which
be rich, burnished gold. The color of Old Forester.
the links between pairs of species map
out what eats what. More abstractly. The golden glo w assures you Old Forester
such webs trace the paths. or food
will taste like no other whisky in the world.
chains. by which energy flows through
the ecosystem. To a very rough first ap That every sip will be mellow, full-bodied
proximation. food webs can be orga and perfectly smooth.
nized into a hierarchy of "trophic lev
els" from primary producers (green What's more, Old Forester gets this color
plants) through herbivores to various and taste solely from the unique way it's matured
categories of carnivores. In the idealized
food web shown on page 160. for exam
in our charred oak barrels.
ple. four trophic levels can be roughly It's made naturally, like Great Wine.
identified. although here. as in most real
food webs. some of the relations do not
But there's another way to test Old Forester's
fit tidily into the classification scheme. superiority that's even more conclusive.
The food web depicted in this exam
ple is typical of ecological systems in the Taste it.
real world. where food chains are char
acteristically short. rarely consisting of
more than four or five trophic levels.
Emphasizing this point. Stuart Pimm
of Texas Tech University has analyzed
data compiled by Joel Cohen of Rocke
feller University for 19 food webs.
which include terrestrial. freshwater
and marine examples. These webs con
tain a total of 102 top predators (ani
mals themselves free from predation). The Great Whisky Made Like Great Wine.
Pimm and Cohen have independently
traced out all the food chains connecting KentuCky Straight Bourbon Whisky. 86 or 100 Proof Brown-Forman Distillers Corp Louisville. Ky 1977.
17 1
i n a l l branches o f b o t a n y , zoology,
o f t a x o n o m i c spec i a l i sts w h o a r e a b l e
to c l a s s i fy p l a n t ( s e e d s , p o l l e n , fi bers,
w o o d , etc . ) , a n i m al ( i n sects, s h e l l s ,
h u m a n r e m a i n s fro m c u rrent
archaeological excavat i o n s . If y o u
y o u r n a m e , a d d ress a n d t h e m at e r i a l s
C u rt W . Beck
V a s s a r C o l lege, B o x 9 2
P o u g h k eepsie, N . Y . 1 260 1
SPEAKER
INFORMATION
KIT.
name
SURROGATE WOODPECKER S, animals that occupy the woodpecker's ecological niche in
c
address
places where woodpeckers are absent, are depicted in this idealized scene. The spe ies shown
include a Galapagos finch, which uses a cactns spine to probe for insects under the bark of
city state zip
trees (0), a Hawaiian bird of the honeycreeper family, which has evolved a .woodpeckerlike
..................... .. .. ..... . . .. beak (b), a striped opossum from New Guinea (c) and the aye-aye, a primate from Madagascar
(t!). Both mammals have evolved long fingers that enable them to extract larvae from trees.
172
environmental stress
crazing phenomenon.
In so doing, they de
veloped a method for
How to squelch
predicting whether or
not poly carbonate will
an unpopular
craze in the presence
of a wide variety of craze
liquids.
The investigation
started with the mea
in plastics_
surement of the criti-
cal, or minimum, strain
at which polycarbonate crazes in numerous liquids.
Attempts to relate these strains to established solubil
ity parameters proved unsatisfactory.
But after conducting experiments with additional
CRAZING MAP OF POLYCARBONATE liquids , our researchers
CrilicaI Slraln (%) di s c ove r e d that for
o Greaterthao 15 liquids having equal sol-
D to 10 1.5
. 0 5 to l0 Ubili'ty parameters, the
!'J Ledan 0 5
measured critical strain
increased with molecu
lar size (liquid molar
volume) .
This finding l e d
them t o incorporate a
molar volume factor
(Vo , on the map) into
the solubility parameter
theory. The result: For the first time, a consistent
ordering of liquids according to critical strain.
General Motors
So how do we now specify the maximum
polycarbonate strain level for a particular chemical
Research Laboratories
W a r re n , M i c h i ga n 48090
environment? The map tells us.
Polymer research: One way of trying to make
environmental stress crazing a thing of the past.
Barcelona Brussels Buenos Aires Frankfurt . Harrisburg Helsinki . s-Hertogenbosch London Luzem Mexico City Montreal
Paris San Juan . Sao Paulo Stockholm Sydney Turin Toromo Tokyo
I I
103 1-
a: Cify
I I
w
CD
::;: State Zip
::>
I I
z
, o Send me bulk rates for my
company/organization.
102 1-
PREVALENCE OF SMALL ANIMALS over big ones is evident in this rough compilation
of the numbers of species of terrestrial animals as a function of the characteristic body length
of their constituent individuals. Disregarding species smaUer than a centimeter in length, which
present difficult problems of classification, the overall trend (broken line) is that a threefold
increase in length corresponds approximately to a tenfold increase in the number of species.
Some representative animals are shown in the illustration, all of them drawn to the sam e scale.
For
er for warm-blooded animals than for derived from rather special assumptions
cold-blooded ones. about community dynamics. The sug-
The conventional explanation of the gestion is. incidentally. representative of
smalI number of trophic levels is that
they are determined by energy flow; if
several recent studies that seek to under-
stand the structure of ecological systems better health,
only 10 percent of the energy entering
one level is effectively transferable to
the Ievel above it. the number of levels is
in terms of the dynamic properties of
the interacting species. Another exam-
pie is the set of studies seeking to eluci-
better health
clearly limited. As Pimm and John Law-
ton 0f the University of York have re-
date the relation between the stability of
an ecosystem (its ability to withstand
information.
cently observed. however. this explana- disturbance) and the complexity of its Improve your health information
tion is not easily reconciled with the food-web structure (the number of spe- and, maybe, your health by sub-
observation that the number of trophic cies and the number of connections scribing to the only health letter
levels is essentially independent of enor- among them). written for the public by the faculty
mous variations in the amount of ener- of a leading medical school.
gy flow and in the transfer efficiencies. he major theme of this article is
In Pimm and Lawton's words, "food T reflected accurately in the discus-
The Harvard Medical School
Health Letter gives you the facts
chains are not noticeably shorter in bar- sion of the length of food chains. Here, about today's health news. Not
ren Arctic and Antarctic terrestrial eco- as elsewhere, the empirical patterns are scare headlines or promises of
systems compared with a productive important, widespread and abundantly miracle cures.
trop lcal savanna or the fish guilds of a documented, but they lack a convincing Every month, you'll receive a
tropical coral reef. " explanation. So it is too for the patterns six-page report covering topics
Pimm and Lawton alternatively sug- that can be discerned in the total num- like: heart attack prevention, the
gest that the explanation may lie in the ber of species, in their relative abun- new pneumonia vaccine, exercise ,
dynamics of the various populations in dances and in the distribution of species megavitamin therapy, the estrogen
the community. They argue, with the among size classes: in place of funda- controversy, cancer, smoking, and
help of mathematical models, that long mental explanations there are only lists more, It's the kind of accurate,
food chains may result in popUlation of possibilities to be explored'. The task straight-forward information you
fluctuations so severe that it is hard for of understanding how ecological sys- need to lead a healthier life,
top predators to persist. This notion is terns work is in the middle of its own Subscribe today,
itself debatable, on the grounds that it is successional process.
175