Janzen, D. H. (1971) - Seed Predation by Animals.

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SEED PREDATION BY ANIMALS 4033

DANIEL H. JANZEN
Department of Biology
University of Chicago, Chicago, Illinois
Access provided by University of New England on 02/13/18. For personal use only.
Annu. Rev. Ecol. Syst. 1971.2:465-492. Downloaded from www.annualreviews.org
Many plants suffer very heavy pre- and/or post-dispersal seed predation
by animals. A few exemplary studies (2,18,38,52,74,87, 106,110,111,124,
140, 161, 162, 171, 181, 187, 196, 197, 203, 205, 207, 217, 220, 227) and a
variety of shorter reports scattered through the agricultural, botanical, and
zoological literature suggest a large and important, yet unexploited, field of
study. It is clear that the pattern of seed predation is highly structured and
that it coevolved at, the chemical, spatial, and temporal level. It involves all
levels of animal-plant interaction from the internal energy budget of indi
viduals to the entire community (203 ) . Owing to parental and sibling com
petition,successful development of a seedling may depend on the seed's dis
persal (101). Equally important,thc seed must escape from the predators at
the seed crop and in the parent plant's habitat before and after dispersal
( 111 ) . The game is played by mobile predators in search of sessile prey;
escape is through a single dispersal move, seed chemistry, parental morphol
ogy and behavior, and evolutionary change.
The processes and patterns are:ideal candidates for ecological and evolu
tionary analyses (83, 111 ) of the typcs traditionally conducted by zoologi
cally oriented bi ol ogists (37, 51,73, 141, 155,156, 182) or applied to plants
in general ( 40,116,208,209,241) .
Seed predators and leaf eaters are often lumped together in ecological
discussions, but they differ in ways especially important in the coevolution
of seed predators and seeds: (a) Like leaves, seeds are highly subdivided
and small,but unlike leaves and other vegetative parts,they have very high
nutrient values per unit volume. ( b) While obvious in large quantities,they
may be extremely inconspicuous once dispersed. ( c) Being comparatively
dormant, seeds have low self-repair abilities but can contain high concentra
tions of toxic compounds (secondary substances) with less sophisticated ad
aptations against self-intoxication than can leaves and meristematic tissues.
( d ) Seed production is not continually required for direct survival of the
parent plant; seed timing, quantity, and quality can therefore be manipu
lated more freely by natural selection than can these traits of leaves and
other vegetative parts. (e ) Seeds are not directly replaced· when killed or
germinated; once removed,seeds may be absent far longer than edible vege
tative parts and thus a common plant species may be rare in time,toa seed
465
466 JANZEN
predator. (f) Once fruits and seeds mature, their death cannot be perceived
by the parent. A major physiological feedback mechanism is thus absent;
the individual plant has only the evolutionary response of its lineage as its
reaction to contemporary qualitative or quantitative changes in predation
intensity.
In the context of this review, animals that eat seeds are termed seed
predators (seed eaters and its more erudite translations leave the fate of the
seed undecided since many dispersal agents swallow seeds, ( 111, 130, 185 )).
With some loss of analytic resolution, I will not distinguish between seed
predators and parasites. By analogy, a bruchid beetle larva eating a legume
seed is as much a parasite or parasitoid as is a chaIcid wasp larva in a moth
pupa; ecologically, the adult bruchid is a seed predator with a daily stomach
capacity equal to her daily fecundity. Squirrels gather food to feed their
young; bruchids put their young on the food. Being a seed predator is a
time-dependent characteristic of the individual; any species' population may
simultaneously contain seed predators, dispersal agents, and neutral mem
bers.
The ecological processes of seed predation apply in many ways to seed
lings, vegetative reproductive structures, and shoot tips, but this comparison
is avoided here owing to space limitations.
Seed predators are restricted to insects, mammals, and birds, and most
species fall in the following families: Curculionidae. Bruchidae,Scolytidae,
Formicidae, Pyralidae, Tortricidae, Olethreutidae, Coreidae, Pyrrhocoridae,
Lygaeidae, Tephritidae, Cecidomyidae, Agromizidae, Torymidae, Eurytomi
dae, Soricidae, Anomaluridae, Sciuridae, Dipodidae, Heteromyidae, Echi
myidae, Muridae, Dasyproctidae, Bovidae, Cervidae, Suidae, Tayasuidae,
Corvidae, Fringillidae, Ploceidae, Colombidae, Phasianidae, Tinamidae,
Psittacidae, Picidae, and various primates. Some members of all plant fami
lies suffer some seed predation, and seed predators are found in all terres
trial and freshwater habitats containing higher plants.
Seed predators and their prey are stilI present in most habitats, but se
lective lumbering and game hunting,pesticides, planting of pure stands, in
troduction of exotic plants and animals, and wild fruit and seed harvest
have grossly and deceptively altered the interaction. The best we can hope
to do is to attempt reconstruction of likely community and population struc
ture from the remnants at hand. What makes this most difficult is that very
often the interaction that selected for the traits still displayed by animal and
plant is now missing, making those traits seem maladaptive or neutral in
significance.
PREDISPERSAL SEED PREDATION

Even though a seed has been released by the parent for later harvest
from the ground by a dispersal agent, its demise is considered predispersal
predation until the seed has been manipulated by a dispersal agent. The in
dividual plant's seed crop is a very important unit of discussion at this
stage. It has size, quality, and proximity traits which may differ strongly
from those of the total seed crop produced by the population or the dis
persed seed crop. On theoretical grounds, predispersal predation by obliga
torily or facultatively host-specific seed predators has a high potential for
operating in a direct density-dependent manner to cause an adult plant pop
ulation's density to exist in a mixed stand at a density lower than would
occur were interspecific competitors the plant population's only challenge
(111). This process should break down most notably where the plant com
munity is sufficiently synchronized and of low diversity to allow seed preda
tors such as squirrels to be highly territorial (203) and to satiate the re
mainder. S imult aneously, the carrying capacity of a habitat for a specific
seed predator may be set by the density of seed crops, irrespective of

whether the seed predator is capable of even approaching 100% predation
on those individual seed crops encountered.
There are many way:; that animals may lower seed crop size before seed
maturation. When a parent plant is defoliated, reduced production of struc
tural tissue should not be the only response (19, 131, 200). Reduction in
seed crop size by defoliation differs importantly from that by nondefoliating
and unprerlictable weather that is inimical to photosynthesis or flower devel
opment (195). The lost leaves maintain an animal population that is likely
to be back for more on a regular basis (leading to directional selection for
defense traits ) , and leaf replacement may exhaust some of the reserves
stored from previous years for seed production. Finally, it may not be "fail
ure" to flower but rather that the plant uses that kind of weather as a cue
not to "turn on."
Direct fl ower eating (35, 45, 62, 136) may lower seed crop size but the
relationship of flower crop size to seed crop size is often rather loose: many
plants produce large numbers of flowers only to attract pollinators, and as a
plant can perceive the loss of a flower, there may be some compensatory
flower production ( 4, 138).
Predation on green fruits or their seeds is common (9, 14, 38, 62, 67, 93,
104, 134, 138, 161-163, 207, 216, 227). Abortion of damaged fruits ( with
resorption?) is a common adaptive response and some compensation of
fruit loss may occur through greater growth of the remaining fruits (128).
Predation on green fruits is very difficult to census since fallen remains are
inconspicuous, rot quickly after falling, and removal from the crop is often
spread over many months. Its los5 is also difficult to evaluate, since the loss
t o the parent in calories and ions must be dependent on the fruit and seeds'
ages. Reserve s that would have been used for fruit maturation may be
stored for later crops (62, 133). On the other hand, coconut palms (Cocos
nucifera ) losing 95% of their green nuts to Pseudotheraptus bugs ( Corei
dae ) increase the number of flowers per inflorescence from 75 to 200, in
crease the number of inflorescences per year from 15-16 to 17-18, and pro
duce morc leaves (226). Animals feeding on green fruits may also intro
duce bacterial and fungal diseases (79,93, 123, 150, 163).
468 . JANZEN
Lacking the space to elaborate the context of the plant and seed predator's
native habitat, to give a straight citation of examples of intensity of predis
persal predation on mature or nearly mature seeds is next to useless. Suffice
it to say that it ranges from less than 1 up to 100% with usual recorded
intensities being between 10 and 90% for animals that are regular seed
predators in their native habitats (13-15,36,49, 50, 55, 59,68, 80, 91, 95,
104, 106, llO, 121-124,126, 127,148, 149,176, 184,192,196, 202,219,238,
244). As discovered by early workers with the biological control of insects,
a severe predator at some prey densities or locations may have little influ
ence on the overall population of its prey if its percent effectiveness is in
sensitive to changes in prey density,and in contrast,a mild predator may be
very effective at prey population regulation. This leads one to conclude

that we badly need descriptions of the sequential relationships of age-spe
cific plant mortality (99) and the effect of these relationships on the produc
tion of new adults by old adults; in this context ·it seems particularly futile
to leave the seed stage out of life tables as is often done (135,231). We
j know almost nothing of the way in which a plant's seed shadow is influ
enced by predispersal seed predation, and we can only make the general pre
diction that it will lower the shadow's overall intensity and that this lessen-
. ing will be proportionally greater at distances far from the parent (111).
Evaluation of predispersal seed predation has numerous pitfalls. X-rays
( 117a,152), opening by hand, flotation, differential staining of crushed dead
seed (28), and rearing (114, 122) have all provided somewhat adequate
means of determining predation intensity but prevent sequential census be
cause the seeds must be collected; seed studies badly need a cheap, light
weight, battery operated, high-resolution X-ray TV camera. Fruits with in
sects in them may open later than intact fruits (244), biasing estimates of
seed predation upwards. Dispersal agents may also select only intact fruits
with the same effect. Further, the seed abandoned by a dispersal agent may
be just as dead as one killed by a seed predator early in the crop's history.
The death of seeds sucked empty by coreid bugs may be attributed to physi
ological seed abortion (126), and unattacked seeds in attacked fruits may
have lowered viability. Entire fruit removal by seed predators is the bane of
seed predation studies; Shaw (196) and others have resorted to censusing
acorn caps. Just as vertebrates that eat fruits ar.e not necessarily seed pred
ators (or dispersal agents ) , insects reared from fruits are not necessarily
seed predators or otherwise detrimental. A large array of insects feed on
seed and fruit fragments after the seed predators have left and the charac
teristic signs (140) of the previous insects may be destroyed.
POSTDISPERSAL SEED PREDATION
The dispersal agents generate a seed shadow around the parent with the
remaining viable seeds. The pattern of a seed shadow's heterogeneity should
strongly influence both the probability that the parent will reproduce itself
one or more times and the detailed location of this reproduction. While
small-scale heterogeneity of the physical and competitive challenge to the
seedling should be haphazard with respect to the location of the parent
plant, seed and seedling predation should be a function of parental·proxim
ity. Further, we may recognize a useful dichotomy among seed predators.
They may be "distance-responsive," with their predation intensity decreas
ing with increasing distance from the. parent tree, or "density-responsive,"
with their predation intensity decreasing with decreasing density of seeds
irrespective of their metric distance from the parent plant ( 111 ) .
We know very little of the distribution of food items sought by postdis�
persal seed predators. The seed shadow should be most intense near the par
ent for wind-dispersed seeds (30, 86) and for some mammal-dispersed seeds
(2 ) , but tropical scatter-hoarding agoutis ( Dasyprocta) may generate

asymmetrical heterogeneity by burying seeds near trees and logs ( 207) and
African duiker (Sylvicapra) may spit seeds into a large pile while chewing
their cud far from the host plant ( 243 ) . Seeds passing through vertebrate
intestines may produce a bewildering array of seed shadow morphologies.
An elephant with 1000 date palm seeds ( Balanites) in its stomach (41 ) is
likely to generate a very patchy seed shadow,just as should mice with many
blackberry (Rubus) and blueberry (Vaccinium) seeds in their intestines
( 5 ) . Taken together, the plants of a �ommunity may generate a very com
plex aggregate seed shadow. At least 72 genera of green or mature seeds
fell or were dropped into 75 seed traps ( 152 X 152 cm ) in one year (aver
age of 7 species per trap ) in a lowland Panamanian forest (207 ) .
A s with predispersal predation, postdispersal seed predation ranges from
very light to very heavy on the community as a whole and for certain spe
cies within the community ( 2, 29, 52, 64, 87, 117, 147, 177, 196,197, 211, 220,
232, 233, 242), but must be viewed in the context of the habitat. Dove
(Z enaida) and other vertebrate predation may be much more intense on
seeds on bare ground than in adjacent grassland ( 18, 52) . Acorns may have
a higher survival rate under leaf litter than in the open ( 197) or buried
more deeply than squirrels (Sciurus) normally do ( 42) ; kangaroo rat (Di
podomys) surface caches may be missed if buried deeply or in exceptionally
dry soil ( 181 ) . Snow-covered conifer seeds have a higher survival rate
since rodents are confined to runways ( 177) .
Even in respect to edaphic conditions and competitive relationships, safe
sites for seeds are easy to conceptualize but extraordinarily difficult to de
scribe and count (100, 101 ) . Predispersal seed predation lowers the fre
quency with which any given safe site is hit by a seed,and if it results in
lowered seed size over evolutionary time-to increase seed crop size ( 110,
203)-it may result in a lowering of the number of safe sites in the habitat
( 111 ) . The probability of a seed being eaten by a predator must also be
taken as a very important characteristic of a safe site. At least it is a mea
surable trait (18,197, 232, 233) .
A plant's seed crop can be easily described and experimented with, but
seed shadows can be very nebulous. Capture of dispersing seeds (30, '
470 JANZEN
207, 222, 230) has the distinct disadvantage of breaking the sequence from
parent to seedling ; radio-tagging (2) probably holds the most promise,
though large seeds may be marked singly (42) or mixed with small ones
that will germinate and disclose the location of a cache (181 ) . Perhaps the
most difficult task lies in excl uding the seed predators with or without ex
cluding the dispersal agents, so as to experimentally examine the relation of
the seed shadow, postdispersal predators, and th e fi nal adult plant density
and spacing; to date, this is feasible, and only partly so, where a plant has
made it to an island but left its seed predators behind (see later section ) .
SEED PREDATOR OR DISPERSAL AGENT?

Whether a seed eater is a predator or a dispersal agent depends in great

part on whether it is after the fruit or the seed, and its precision in seed
processing. Packrats (Neotoma) eat entire prickly pear fruits (Opuntin)
and the seeds pass through unharmed; the same animal eats the outsides of
some species of cholla fruits (Opun.tia)
other cholla species, it husks the seeds and chews them up ( 210) . Gallina
ceous birds may use hard seeds as substitutes for grit (20) and their fate is
probably to be ground until dead. Bird seed predators can be very deadly; of
40,025 seeds of at least 20 species fed to linnets (Aca.nthis), only 7 made it
through the birds and germinated ( 185) . Since many seeds are poisonous to
vertebrates (see later section) we may expect such birds to be careful about
what they eat, especiaIly since they may eat a large number before the giz
zard grinds through the seed coat. The linnets mentioned above rejected
seeds of at least 9 species on the basis of taste, and some are seeds known to
contain nitrogenous toxins (Medicago, Convolvulus, Papaver); doves and
other birds may do the same with toxic seeds ( 52, 130) . While chewing
their cud, duiker may spit out part of the seeds they have eaten ( 243) , but
that is not necessarily the fate of all seeds eaten. While some seeds have
" improved germination" after passage through a vertebrate gut ( 130)
( though the proper terminology is probably that they have "retarded germi
nation" if they do not pass through a vertebrate gut), this by no means
indicates that all seeds eaten by an animal make a safe passage. Some large
tropical birds regurgitate some of the large soft seeds in the fruits they eat.
The timing of this act, its physiological induction, and its adaptive signifi
cance to the bird are uninvestigated; to the plant it may be the only way to
safely disperse a large soft seed, and to the bird it may be a way of avoiding
seed toxins and of rapidly emptying the stomach for more fruit. Seeds "in
tended" for a dispersal agent may well be killed by another, such as when
starving squirrels prey on seeds of Rosa and Arctostaphylos ( 202) , which
are normally bird dispersed. Seeds rejected by harvester ants after capture
( 94) or lost by kangaroo rats ( 2 1 8) around nest entrances may survive to
adult status and may even be growing on nitrogen-rich soil owing to their
proximity to the nest.
When an animal is regularly the seed dispersal agent and seed predator
for the same plant species-as mice, squi rrels, and agoutis are to large
seeded trees (2, 42, 196, 207) and as kangaroo rats may be to small-seeded
desert plants (181) -we must recognize the seed predation as the cost of
reliable dispersal and directly analogous to a juicy fruit or complex explod
ing pod. The cost may seem high: gray and fox squirrels recovered 415 out
of 419 buried hickory nnts (Carya) during a Michigan winter (42), mice
and other rodents recovered 86 out of 92 white pine seed caches during a
Massachusetts winter and spring (2), kangaroo rats (Dipodomys) re
covered 83 to 85% o f surface seed caches at their normal burial depth
(181), and agoutis probably recover most of the seeds they bury (207).
However, many of the surviving seeds have been moved away from the par
ent and nicely buried. ( Perhaps there has been a coevolution of the distance
under ground that a rodent can smell seeds and the distance that a seed
should be under ground for moisture purposes. ) This is probably a much
higher score on hitting safe sites than could be achieved through inanimate
dispersal with a large seed. Formosov (76) feels that Pinus cembra seed
burial by Siberian nutcrackers (Nucifraga) is responsibl e for most popula
tion recruitment by the pine population. The effect of squirrels caching
cones in wet sites to prevent cone opening (wi nd -di spersed seeds ) mayan
first sight appear similar, especially since they may drop one seed for every
one eaten; however, on account of the squirrel's activity and cone debris
piled on the seeds, it is probably a very poor site. The seed-burying animal
may not be the only predator on seed caches; Sitophilus granarius (Curcu
lionidae) is a common pest of Old World granaries and chose shelled
acorns out of a variety of othe r seeds and grain to feed on (107).
Relative food abundance may have a strong effect on whether a verte
brate is a predator or dispersal agent. I cannot agree with Smythe' s (207)
evaluation of this aspect of dispersal when he states that "the efficiency of
dispersal depends on the number of seeds eaten, and if at one time of the
year there are a greater number of seeds available than can be eaten, then
the energy that went into the production of the surplus will be wasted."
\\That he probably meant was that the efficiency of dispersal depends on the
number of seeds carried off and not eventually eaten, and if at one time of
the year there are a greater number of seeds available than the entire com
munity of animals can kill and/or carry off, and if there is no chance of a
seedling sur viving below a parent, then the surplus is wasted.
CHEMICAL ASPECTS OF HOST SPECIFICITY

Seeds of plants, and especially those of large plants, are notorious for
their diversity and hi gh concentrations o f "secondary compounds" toxic to
some vertebrates, insects, or microbes. These compounds are not likely to be
waste products (110, 118, 241), and many seeds toxic to some animals are
not toxic to others. These observations are compatible with the concept that
seed chemistry has coevolved with seed predator host specificity. Strong
obligato ry or facult ative host specificity by the seed predator is a key requi-
472 JANZEN
site to its act o f i nfluencing the density and spacing of adult plants on the
one hand, and being limited by the seed crop resource on the other hand.
This specificity is also strongly influenced by fluctuations in prey availabil
ity, which will be discussed in the following section.
Host specificity should be seen from the plant's viewpoint as well as
from the animal's and in the context of its native habitat. If a Tamiasciurus
squirre1lives primarily on the cached cones of white spruce (Picea glauca)
and only eats large amounts of other items (e .g., P. mariana) when forced
(34, 124, 205), it is a highly host specific seed predator. A species of Curcu
lio with a major part of its br eeding population in the acorns of each of two
sympatric species of oak (92) is a generalist from the viewpoint of either
oak,but a specialist from the "Quercus viewpoint." Even an insect that preys
on the seeds of ten species of tree can be an extreme specialist if the crops
are allochronic or exist in a tropical rainforest of 500 tree species.
There is no reason to believe that the chemistry of fruit and seed loca
tion by seed predators differs in any substantial way from that of other ani
mals feeding on plants. It should therefore involve attractants, intoxicants,
deterrents, sign stimuli, and feeding stimulants ( 108, 132, 153, 246); care
should be taken to distinguish between attractants adaptive in that sense
(e.g., sugars in fruits) and attractants with negative fitness (those com
pounds that the animals have come to recognize over evolutionary time as
signaling the presence of prey). The chain of sequential chemical cues may
also be short-circuited with seed p redators (31, 56), as with other plant eat
ers,and this leads to oviposition or feeding on new "hosts" in the laboratoryor
agricultural area.
Seed colors are probably generally adaptive in respect to visually orienting
seed predators, as is so well illustrated by mourning doves differentiating be
tween gray (toxic) and cryptic (palatable) seeds from the same dove weed
( Eremocarpus setigerus) plant (52). Agoutis may move as far as 50 feet
toward the sound of a falling fruit (207), and there probably is strong se
l ection for l ar ge nuts to have weak odors so that,once buried,they are diffi
cult to locate.
The formation of a search image, or habituation to a particular seed,
should be extremely impor tant with facultatively host specific seed preda
tors (1 02, 219, 220). That kangaroo rat (Dipodomys) seed caches o ften
conta in large amounts of one species of seed (218,229), the repeated visita
tion of the same tree by British bullfinches ( PyrrhuZa) (162),and the total
stripping of the seeds from one desert annual by Veromessor harvester ants
(220) may be manifestations of this. Since the bullfinches returned year
after year to the same individual trees,there is a suggestion of exception
ally strong selection against trees weak in defensive compounds.
Fruit clwracteristics ..oC-The primary adaptive functions of fruit morphol

.
ogy, chemistry, and behavior are seed protection and dispersal to safe sites.
In respect to a specific predator, very fine details of fruit morphology and
chemistry may be highly adaptive (llO)-examples are husk hardness (7,
28a, 150,203), rates of husk expansion (75a), hairiness ( 147, 238a), con
strictions separating seeds (75a), liquid resins ( 106, 184), and direct intoxi
cants like gossypol (238a). Once a seed is mature, the timing of the adver
tisement of this and its release should be influenced by seed predators as
well as dispersal agents and seasons for germination. This is particularly
obvious with wind-dispersed conifer cones; the race between how fast
squirrels can cut and store them and how fast the seeds are shed (205,219)
has had a strong coevolutionary component (203) . Were a Tamiasciurus
red squirrel to harvest seeds off the ground individually, it would have to find
one every 10 to 15 seconds all day long ( 205). Black spruce cones (Picea
maritima) are only rarely harvested by squirrels ( 34, 205 ) and they dis
charge their seeds gradually (5) .
We may expect animals to be continually fine-tuning their behavior to
these plant traits, as well as vice versa. Red squirrels in areas with large
amounts of tough serotinous cones (Pinus cont,orta) have better developed
-
jaw mechanics than those in areas of softer cones (e.g., Douglas fir) (203).
There is greater variance in female Curculio proboscis lengths (used to pre
pare oviposition sites in the fruit wall or through it, as well as to feed) than
in those of males ( 92).
Seed co at While hard seed coats may keep out a large number of gen
.-
eralists, they are probably not as important as internal seed chemistry in

influencing predation. Palm seeds, encased in a bony endocarp and charac
teristically lacking internal toxicity, are probably exceptions. In Hawaii, the
Itamarca strain of mango (Mangifera indica) has a seed too hard for first
instar larvae of Sternochetus mangiferae (Curculionidae) to drill throug h ,
and they wander about in the fruit and die. S. gravis of Burma eats only
the mango fruit (14), which may be the evolutionary consequence of this
kind of resistance by the plant. Pecans ( C ary a) with cru shing strengths of
66 to 79 pounds are ground through by t urkey gizzards in ab out 1 hour,
while hickory nuts (Carya) with crushing strengths of 167 to 257 pounds
require 30 or more hours (191). Th ose acorns that germinate in the fall are
very susceptible to attack by C onotrachelus weevils Dvipositing through the
split seed coat ( fruit) wall (91 ) . Seed coats may be toxic in some cases;
doves reject seeds by taste (52), but this could be a simple case of associat
ing a distinctive taste with a later unpleasant consequence. Endrin-coated
acorns do not poison gray squirrels (S ciurus carolinensis) because they
shuck them, but they do poison cotton rats (Sigmodon hisPidus) because
they gnaw through the coat (115) . The resin in fir seed coats (Abies) may
deter squirrels and mice (203).
Internal seed chemistry .-D irect toxicity may be based on 1 to 10% or

more concentrations of such things as selenium ( 10, 223), cyanogenic glyco
sides ( 1 18), saponins ( 98), g ossypol (139), a t rem endou s variety of alka-
474 JANZEN
loids and uncommon amino acids (21,77,110,113,221,240) ,and endopepti
dase inhibitors ( 8). Where space ( = weight = food reserves) is a limiting
resource, as in seeds, it would not be surprising to find selection has favored
the use of toxic compounds that are simultaneously storage products,
though certainly some are not metabolized by the seedling (224; E. A. Bell,
personal communication). Inert fillers, such as silica (66) and cellulose,
commonly function as defensive compounds in vegetative tissue and are
generally absent from seeds. There is no compelling reason for the defen
sive compounds in a plant's seeds to be the same as those in its vegetative
parts, since the challenging animals are likely to be quite different. The cy
anogenic glycoside polymorphism in Lotus foliage is missing from its seeds
(118) , and while the joints of cholla and prickly pear cactus ( Opuntia spp.)
are eaten avidly throughout the year by Neotoma packrats, there is strong
differential feeding by packrats on the fruits and seeds of these two species
(210) . On the other hand, a good general poison selected for by one animal
is then subject to selection for its general distribution throughout the plant.
Mimosine is toxic to browsing mammals but found throughout Leucaena
glauca plants (221); bruchids and an anthribid that attack the seeds are
resistant to it ( 106,110 ) , but general seed predators probably are not.
This brings to mind the obvious point that a seed toxic to most animals
in the habitat may well have specialists that can feed on it. Particular spe
cies of bruchids have evolved resistance to the toxic alkaloids and free
amino acids in seeds o f Erythrina (33) , Astragalus (114) , Abrus (32) ,
Dioclea ( 113) , and Sarothamnus (171) , and boll weevil attraction to gossy
pol in cotton ( 139) provides an example from the Curculionidae. We must
recognize, however, that the actual evolution may be taking place in great
part at the level of the microflora in the seed predator's intestine. If the
seed predator is small relative to the seed, then we may expect a gut with
one or two species of bacterial specialists that vary between species of seed
predators. If the animal is large relative to the seed, then we may expect a
large microflora community which would be severely disrupted by a high
concentration of any one species of seed. Ruminant artiodactyls can have
their intestinal flora severely disrupted by high concentrations of plants
normally eaten in small amounts (158, 173, 199) , and seed predators also
should ( gossypol in cotton seed is toxic to nonruminating animals, 139) .
Squirrels apparently can eat small amounts of canavanine-containing seed,
but large doses are probably toxic (113) . It is interesting that the two big
groups of nonruminating artiodactyls, Suidae and Tayasuidae, feed in great
part on roots and on seeds involved in predator satiation rather than those
that escape by being toxic. Since many toxins are antimetabolites whose
effect can be reversed by high concentrations of other specific compounds
(77) , we may expect ability to eat small absolute amounts of some seeds to
be a function of animal body weight. Further, the predator's developmental
stage should be of importance; the older a chicken, the less susceptible it is
per unit body weight to the toxic amino acids in Lathyrus seeds ( 179) .
Beech nuts, apparently totally lacking in toxic compounds, are found signifi
cantly more often than acorns and other seeds in the stomachs of juvenile
fox and gray squirrels than in adults ( 166 ) . While lygaeid bugs can feed on
a va r iety of plants to sustain maintenance metabolism, they require very
specifi c seeds for reproduction ( 74 ), and this may be a function of secon
dary compounds as well as of conventional nutritional traits. After all, the
cost of neutralizing a toxin must be subtracted from the nutritional value
gained, even if a seed predator can eat a toxic seed.
Direct feeding trials with seeds can bring out very obscure seed traits
( 31, 34, 52, 59, 60, 61, 74, 105, 126, 174, 203 ) , but even here it is often very
difficult to identify the actual toxic agent. Now that several seed predators
may be reared on partly or completely defined diets ( 105,183, 214, 215 ) , at

least suspected compounds can be directly tested. As Eyles (74) has so dra
matically shown with lygaeid bugs, in such tests "toxicity" must not be
view ed as an absolute trait but rather a relativ e one, affecting growth rates
and fecundity as wcll as mortality rates. Observations on seed predator se
lectivity in the field are grossly complicated by the relative abundance of
seeds, as shown by the difficulty in interpreting the seed species composition
of kangaroo rat seed caches ( 181) and bud and fruit preferences of finches
( 161, 162 ) .
Lack of toxicity.-Plants involved i n population and communitywide

-
predator satiation ( usually temperate-zone trees, but perhaps southeast

Asian Dipterocarpaceae as well, 111, 203 ) appear to have seeds edible to a
very large variety of seed predators. Practically the entire vertebrate com
munity of the Russian taiga feeds on Pinus cembra seeds (76, 217 ) , and at
least 44 species of mammals and 37 species of birds eat North American
coni fer seeds (204 ) . Lepidoptera larvae infesting conifer cones tend to have
very long natural host lists ( 48, 54, 129, 140 ) . Fagaceae ( acorns, beech nuts,
chestnuts) are eaten by a vast host of vertebrates and insects ( 67, 91, 92,
145, 157, 196, 206, 233, 235 ) , though their predators show some detailed
specificity which may be based on husk traits (36, 91) or on seed toxicity
itself ( 71, 233 ) . This apparent lack of toxicity is probably associated with
the general phenomenon that plants escaping through predator satiation ap
pear to put comparatively little effort into direct internal defense of seed
( 110 ) .
The general lack o f toxicity o f grass seeds exemplifies a n important
aspect of the interaction between seed toxicity and predator satiation. Being
individually small and not enclosed in a large fruit, grass seeds are difficult
prey for large predators, except those that can harvest each individually
( harvester ants, birds, kangaroo rats, mice ) . These animals, however, re
quire large numbers of seeds. Each grass plant produces only a very small
number of seeds, and there are a very large number of grass plants in most
grass communities. The chance of any in dividual predator encountering
several seeds sequentially from the same plant is low. The seeds of a mutant
476 JANZEN
strain of grass would have to be exceptionally toxic (and therefore be se
verely limited in nutrient reserves) to cause a predator to selectively avoid
them, and the predator would have to be incredibly perceptive to remember
to associate with the toxic effect a few nondescript seeds out of the thou
sands it eats. With a tree, however, the large seed crop is likely to provide a
major part of the predator's diet for many days, and thus a mutant with
toxic seeds is likely to have lowered predation.
Caloric content.-Some vertebrates may harvest seeds largely as a func

tion of caloric content (120, 143, 203), but where toxic compounds or diffi
culty of husking confuse the picture (161, 162, 187), the relationship may
break down. Not only must the cost of extracting ( 187) and catching the
seed be considered, but the actual caloric value of a seed to a seed predator
may be much lower than that recorded in the calorimeter ( 125 ) . A plant's
Latin name is not a certificate of quality; cones from fertilized trees may be
more heavily attacked by squirrels and insects (89,202), and 144 new white
spruce cones (Picea glauca) per day will support a red squirrel whereas 194
old ones are required (205). The smaller the vetch seed (Vicia), the
smaller the resultant bruchid (65 ). Even ·Conotrachelus nemtphar weevils
reared from different commercial fruits have different fecundities and dia
pause behavior (146).
This brings up the point that a common cause of host shifting of wild
seed predators to crop plants (9, 22, 46, 132, 162) is not only that the crop
plants are grown in pure stands (81, 112), but that the natural toxins have
been bred out of the seeds (96)-and replaced by pesticides (112).
Variance in susceptibility.-Entirely from data on agricultural plants

( 11, 12, 13, 67, 69, 139, 164, 213 ), we can state that there may be strong
interpopulation variance in susceptibility to seed predators, which presum
ably reflects an original intrapopulation variance that the predator acted
upon. Within one seed crop of Mucuna, L-dopa concentration ranged from
6 to 9% dry weight per seed (21). Intertree variance in ash seed oil contcnt
may be responsible for selectivity shown by British finches (162) and there
is considerable variation in bruchid survival in some species of commer
cially grown seeds (105).
PREDATOR SATIATION
Predator satiation is a highly coevolved interaction of parental behavior
with seed predator behavior (72, 110, 189, 203, 217), just like direct fruit
and seed traits, and the plant population cannot be treated as a resource
base unchanging over· evolutionary time (133). There are four critical as
pects of predator satiation: all seeds of a seed crop (of a fruit, tree, popula
tion, or community) are not equally available to all members of the seed
predator population, the seeds are available for only a short time, the preda
tors are partially to totally host specific to the seeds in question, and the
predators cannot maintain high pop ulation densities in the prolonged ab
sence of seeds.
Within the individual plant's fruit cr ap . At this level of integration, the
-
important variable is how fast the plant can mature and disperse its seeds
(110, 196) , as compared with how fast the individual seed predators in its
neighborhood can find the crop. Seed crop size is a critical trait and it is
probably adjusted over evolutionary time through changes in seed size (101,
1 1 0, 203) and through changes in the length of time between crops ( see
later s ecti ons ) In general, within a major taxonomic group and within
.
broad habitat types, those plants that escape by predator satiation should
produce larger amounts of smaller seeds in any given seed crop than those
that escape through direct chemical defenses (110 ) . Selection for larger
seed crops should also select for gre ater age at first reproduction (1 1 1 ) , and
with age there may even be a local buildup of predators on an individual
tree's seed crop (119) . Since oak seed crops may increase and then decline
with age (70 ) , predato r satiation may be maximal at an intermediate age of
reproduction. The maximum destructive ability of the individual animals
that encounter a seed crop is poorly known, but ranges from values like 15
eggs a day (equal to a maximum of 15 seeds a day) and 100 to 300 in three
months for Sternachetus mango weevils (7,9, 14) ,50 to 100 eggs in a life
time for bruchids (110, 132, 225 ) , 537 sequoia cones cut and stored in three
days by a Tamiasciurus squirrel (198) , 97 to 137 white pine seeds eaten per
day by mice (1) , and 100 acorns per day by Engl i sh wood pigeons (196) .
This is strongly complicated when a seed is slightly toxic so that the number
that can be eaten per unit time is considerably lower than the actual stomach
capacity (113) . The seed predators may build up on a seed crop once it is
located, and thus the probability of a seed surviving becomes age dependent.
However, this may be disrupted by the need for many insect adults to leave
the host plant and seek other types of food before ovipositing ( 114, 165) ;
if seed crops are close together, as in temperate zone communities, this may
only result in considerable exchange of insects between crops; but in tropical
communities where trees of the same species are far apart, it may result in
considerable loss of seed predators.
Seed predators move widely varying distances in locating individual
plants in fruit: extremes range from 10 to 130 foot foraging ranges of har
vester ants (211, 220) to a 4 hectare area for a red squirrel (124) to perhaps
several miles for parrots. Even within a tree's crown there may be differ
ences in intensity of satiation (17, 129) , underscoring the general problem
that distances between seed crops have to be measured in the percent of
seed predators that succeed in crossing,not in metric units (111).
During any given fruiting season, the timing of an individual's seed crop
may be influenced (through selection ) by the presence in the community of
other species' seed crops (109, 110, 111,203) ; this may occur both through
consecutive exchange of seed predators in common and by having the pred
ator attracted away from one species' crop by another.
478 JANZEN
The more difficult it is for predators to move between crops, the more
effective should be predator satiation at the individual crop level (36, 50,
106, 1 1 1, 140, 239) . This assumes that pollination is not the limiting factor
as plants become more widely sepa rated and that the l ong interplant dis
tance is not simply a re flection of poor site quality that in turn is expr essed
as smaller seed crops. However, in pure stands of one species, decreased
tree density may lead to decreas ing l y effective pr eda tor satiation (129, 167,
190), as t he predator may still move easily between crops but there a re
fewer seeds pe r a cr e.
At the population and community level.-Predat or satiation is undoubt

edly responsible for the spectacular coevolutionary (somewhat cyclical)

fluctuations in seed predators and their carnivorous predators in forests
across the upper half of North America and Europe ( 133, 203, 217) . The
system probably developed through the following general evolutionary pro
cess ( 133, 203): A weather event either destroys a flower crop or fails to
provide a flowering Cue one year. The sterile individuals have a larger crop
the next year and satiate the p red ators more effectively, leading to selection
for those plants hypersensitive or hyposensitive to the disruptive weather
ev ent. Further, there is selection for an internal physiological system pro
gressively more responsive the longer the tr ee h as remained sterile ( = the
more food reserves it h as) . The external weather events must be of suffi
cient intensity to override local differences, such as those fou!ld in compar
i ng a dry ridgetop with the adj acent riverbottom, and can be badly confused
by clearing la nd for agr icultu re . While the eventual trait is recorded as a
"population trait, " it is clear that the population is really not synchronized,
but rather that the individuals are synchronized with weather events that
symbolize low seed predation. It is clearly incorrect to speak of crop "fail
ures" in contemporary communities and to treat peak seed production years
as "normal." For any given species of tree, the constraints to l ength eni ng
the time period between crops-2 to 10 or more years ( 16, 53, 55, 78, 88,
133, 203, 217)-will be set by the comp etitive disadvantage of not putting
seeds into the habitat when available, the costs of storing nutrients, and the
proba bility that the individual wiII be heavily damaged by fire or insects
b efore frui ting. As the individuals of a major tree species in the community
become synchronized, selection should build up against those asynchronized
species to ei ther drop out of the community or become synchronized, which
brings in the complication of the coexistence of species that take 1 versus 2
years to mature fruits from ,flowers (78, 203) .
Predator populations have two crit ical functions in this system. First,
they are likely to produce a local buildup in density on a peak seed-produc
tion year which may be reflected in decreased territory size ( 1 24), de
creased aggressiveness between adults and juveniles (206, 235), increased
bree ding ( 76, 78, 85, 88, 133, 203, 217), and mass immigrations ( 144, 145,
157, 196, 203) ; we can no longer study the best examples of the latter case
in North America, owing to the extinction of the passenger pigeon and the
Carolina parakeet.
Second, the predators generate intense selection against those individu
als fruiting out of ph ase ( 55, 70, 78, 90, 97, 129, 137, 140, 166, 172, 180, 190,
203, 219, 233, 238 ) . There may be a complicating polymorphism in the case
of insects that diapause between peak years and in that of birds that move
long distances to large seed crops. Shaw ( 196) points out that while a
heavy acorn crop produced twice as many seedlings per square meter than
one a fourth as large, the light one produced nearly three times as many
seedlings per acorn. An explanation of this paradox is that the light crop was
not big enough to attract flocks of wood pigeons ( Columba ) , a major preda
tor ; we may expect that an intermediate-sized crop might have attracted the
birds, but not sa tiated them.
The seed predator is confronted with the problem of waiting out the
time between crops. A variable fraction of the insect population generally
goes into total diapause for 1 to 5 years ( 36, 55, 92, 103, 104, 121, 129, 140,
228) , a behavior strongly reinforced by severe competition among insects
for the cones in off years ( 129, 140). Where diapause is highly synchro
nized with the plant population, percent seed predation on the peak years
may even be higher than during the off year ( 58, 129 ) . Synchronization of
the insect with the plant population may be particularly difficult with those
plants that initiate flower development by a cue occurring 2 years prior to
seed maturation, as occurs with some oaks and conifers ; even this timing
may have its adaptive significance solely in increased predator satiation.
Cache hoarding, associated territoriality ( 124, 142, 203 ) , and migration
(217) may all be viewed as analogous adaptations by vertebrates to waiting
out the off years. Scatter-hoarding and overlapping home ranges (25, 188,
207) may be viewed as analogous to going into reproductive diapause, as do
some weevils (9, 14, 146) . Either in diapause or as active adults, waiting
between seed crops in tropical habitats should be particularly difficult since
poikilotherm predators on insects do not cease their search until they run
out of prey ( in contrast to what occurs during northern winters) and the
seed predator will undoubtedly be burning up more reserves ( = lowered
fecundity) than its counterpart waiting in a cold climate.
In tropical communities with high tree species diversity, the shortage of
dramatic synchronization cues, coupled with the lack of a population large
enough to satiate the specialists and generalists, makes synchrony of entire
populations or communities unlikely, though synchrony may be approached
as seasonality increases ( 111, 207). Satiation thus has to be primarily at the
individual level and may differ somewhat from satiation achieved through
synchrony with other members of the population or community. The iso
lated tropical tree with an annual seed crop is likely to have its own local
population of insects specializing on its seed crop ( e.g., mango weevils on
Mangifera, 14 ) . If a mutant appears that waits long enough between crops
so that the local population is severely depleted, the only density-dependent
480 JANZEN
seed predators will be those that arrive at each crop through immigration,
and the consequences are sel f-evident. This may even occur within a year,
as when Lygus bugs fail to build up in years when weather causes two dis
tinct flowering peaks by African coffee bushes, but do heavy damage when
there is continuous flowering ( 1 36 ) .
A core aspect o f predator satiation i n high species diversity communities
is increased individual seed crop size, and we may expect this to generate
two kinds of dioecism. Obligatory dioecism may be selected for by any
adaptive value of increasing female seed set over that of the hermaphrodite
( 186). A byproduct is that to the seed predator the trees in a dioecious pop
ulation are rarer than they are to the forester ( 1 1 1 ) . Further, where there
is strong overlap of generations, a female tree that adds half males and half
females to her vicinity will have a lower rate of mutual exchange of seed
predators with her offspring than will a hermaphrodite producing hermaph
rodites. There are substantially more dioecious tree species in lowland tropi
cal communities than in their temperate zone counterparts ( I ll ) . Faculta
tive dioecism may appear through a tree on an edaphically or competitively
good site, satiating p redators at the individual or community level, while it
may be more advantageous for its less fortunate sibling only to "capture" as
many ovules on neighboring plants as possible by means of pollination. Cer
tain individual oaks fruit consistently on peak years and others never have
large crops (47, 195) ; acorn yields are higher from trees with spreading
crowns (58), and individual coni fer yields are increased by thinning ( 6 ) .
COMPETITION BETWEEN SEED PREDATORS

Competitive encounters between seed p redators should be frequent, ow
ing to the finite and often small quantity of their h igh-quality food. Where
seeds escape primarily through seed toxicity, this should be expressed over
evolutionary time by such things as character displacement of crop use (two
Acanthoscelid es bruchids on the same As tragalus population having differ
ent times of emergence, 1 14) or host specificity ; I have found almost all o f
at least 40 species o f bruchids in a Costa Rican deciduous forest t o o ccur on
no more than one host. This type o f displacement should also lead to strong
divergence of the chemicals and other plant traits that mediate host specific
ity. Where seed escape i s primarily through predator satiation of a more
generalized community o f seed predators, we may expect very stable territo
riality where prey species diversity is very low ( 124) ; this progresses to
more and more overlap o f home ranges and scramble competition ( coupled
with mass "migrations" on "off" years ) as the species diversity of the forest
rises (and the probability of a territory always containing some trees in
fruit declines )(3, 124, 203). Some specialization may occur here as well
(43, 234). For different groups of predators in a given forest, the situation
may be different ; to the mice and chipmunks living on herb, shrub, and con
i fer seeds, it may be a much more diverse community than to the red squir
rels dependent on two to four conifers for reproduction. This is especially
evident in the dynamics of Apodemus and Clethrionomys mouse populations
in European oak forests (206, 234, 235).
For insects, there may be direct fighting for ownership of the inside of
a fruit or seed (23, 27, 55, 106, 1 29, 140, 1 71, 244) resulting in cannibalism
( 5 5 ) , nearly complete population diapause between seed years (140 ) , and
sometimes, restriction of otherwise generalist species to a single host ( 1 75 ) .
Lowered mortality from competition, plus a large number of insects coming
out of diapause, may lead to an increase in absolute seed mortality on peak
crop years ( 1 29, 1 40). Over evolutionary time it is probably in part respon
sible for the very uniform distribution of the species in the large genus Cur
culio over the species of North American Fagaceae ( 92).
The density of seed predator populations and communities seems gener

ally to be set by the sizes, frequencies (in time and space) , and quality of
seed crops ( 97, 124, 133, 1 57, 161, 162, 203, 217, 235 ) . The actual density
dependent mechanisms are the usual proximal causes, such as increased so
cial aggression as food gets scarce, starvation on suboptimal foods, in
creased susceptibility to weather and predators by migrating or weakened
individuals, etc. Only in one case has it been shown that with the removal of
a specific carnivore, the density of a seed predator rises ( 39), and here the
carnivore's removal appears simply to temporarily raise the carrying capac
ity of the habitat for the seed predator.
PARASITES AND PREDATORS OF SEED PREDATORS
The insects preying on large temperate-zone seed crops characteristi

cally support large parasite populations (23, 24, 63, 1 14, 121, 159, 160, 169,
171), as do the bruchids from very common tropical hosts such as A cacia
farnesiana. There is, however, almost no evidence of entomophagous para
sites of tropical host specific seed predators ; this is not surprising since
their trophic position is that of a hyperparasite in a classical parasite-host
relationship. Even in temperate-zone forests, it is commonplace for an ento
mophagous parasite of a seed predator to attack a wide array of host spe
cies ( 24, 57, 236) , and Cerceris wasps are restricted to one higher taxon
such as Curculio ( 193 ) , Bruchidae ( 237 ) , and Tychius (212).
One can assume that both insects and vertebrates moving between seed
crops are subject to normal generalized predation, and some vertebrates
may prey heavily on insects while in the seed or fruit (36, 59a, 67, 90, 122 ,
168). Owing to the weakened seed coat, most seeds containing insects are
probably digested when eaten whole by dispersal agents, but this need not
always be the case (151).
SEED PREDATORS AND FOREST COMPOSITION

While there are strong logical and circumstantial reasons to suspect that
seed predators have a strong influence on the apportionment of the total
plant community biomass among the plant species present (26, 1 1 1, 203 ) ,
there are almost n o experimental field tests. Exclosure experiments (18, 75,
482 JANZEN
1 97, 232, 233 ) have been conducted on a very small scale or in respect to
only one species. Rodents and harvester ants may influence the rates and
directions of plant succession (26, 154, 1 80, 181, 194, 220 ) . In favoring the
seeds of one major plant species over those of others, even apparent gener
alists may have a strong effect (220 ) . It is probably not a coincidence that
in a mature Ohio forest where less than 15% of the reproductive trees were
hickory and beech (oak was 67% ) , more than 50% of the seed diet of fox
and gray squirrels came from these two species (166 ) . There is strong fa
voritism for certain conifers in predispersal seed predation by squirrels
(203) , and postdispersal seed predators left 12, 31, and 65% seed survival
of Douglas-fir, western hemlock, and red cedar in the same Oregon c1earcut
(84 ) . This should influence stand density, but there is no reason why pro
portions of adult trees should match up with intensity of predation. A tree
with very competitive seedlings may suffer strong predation yet be more
abundant, or of the same density, than one with weak seedlings and light
seed predation ( 1 1 1 ) . We can only state with certainty that if seed preda
tors are removed over contemporary or evolutionary time, the relative
abundances of specific plants are likely to change.
In general, we should expect that as a higher proportion of the members
of the plant community escape from seed predators through population and
communitywide predator satiation ( rather than primarily through chemical
means and great interplant distances ) , the species diversity of the commu
nity should decline. This is so because the community structure will be set
progressively more by interspecific plant competition alone. In short, from
the aseasonal tropics to the seasonal tropics and the temperate zones, the
seed predators probably become less and less effective at keeping any partic
ular tree species from competitively excluding all other species with a simi
lar life form ( 1 1 1 ) . The epitome is probably the Rttssian taiga, where the
density of Pinus cembra is probably set only by intraspecific competition ; it
is interesting that its 8 to 1 1 year periods between seed crops ( 76, 217) are
among the longest for a conifer. The exceptions should lie in instances
where a chemical defense is so effective that nothing can eat the seeds,
which is probably the case with tropical mangrove swamps of notoriously
low tree species diversity, or with islands where seed predators have not
become established.
SEED PREDATORS ON ISLANDS

Owing to the periodic nature of seed crops and to the low absolute num
ber of habitat types available on islands, we may expect establishment
and persistence of seed predator populations on islands to be rare (208) .
Even very large islands may have depauperate seed predator communities,
as evidenced by the absence of Curculio weevils and other insects from the
acorns of British oaks ( 196 ) and the replacement of the British red squirrel
( Sciurus vulgaris ) by the North American gray squirrel (S. carolinensis)
during the past century ( 138a) . The manifestation of a depauperate seed
predator community should be pure stands of species of plants which a re
much rarer on the mainland and lower species diversity than would be ex
pected on the grounds of interspecific plant competition alone. It is notewor
thy in this connection that "the greater part of the natural forest cover [of
Hawaii] is composed of a relatively few plant species," and only the larvae
of a tortricid moth attack the pods of Acacia koa, one of the most abundant
tree species (82) Pseudotheraptus coreid bugs are present on some islands
.
off Zanzibar ; there coconut yi elds are 5 to 10% of those on i slands where the
bugs are absent ( 226 ) . This may be a self-reinforcing system whereby those
plants that become established on an island occupy the habitats so thoroughly
( owing to lack of a seed predator ) that other plants find it much more diffi
cult tobecome established than they would if the occupants were at the den
sity maintained on the ma i nland .
Plants introduced onto islands are notorious for producing large pure
stands, and there is no reason to believe that this is entirely due to release
from competition. When seed predators are later introduced, they may take
a heavy toll of seeds ( 106, 123, 149, 201), but if at the time there are no
other plants to compete with the plant under attack, it may be some time
before another encroaches on its habitat.
We may e xpect some extremes of predator-prey interactions to occur on
islands, as with the huge slow-growing fruits ( and tiny seed crops) of Lo
doicea maldivica palms which occur in pure stands on the Seychelles is
lands, which are presumably free o f indigenous seed predators. A strain of
wild Mexican cotton ( Gossypium ) from an island in the Gulf of Cal i fornia
has 3.5% gossypol in the seeds, as contrasted to 1.2% for mainland strains
(139), probably indicating the presence of a well-established cotton seed or
fruit predator. On small Caribbean islands, fig trees (Ficus ) have strong
intratree fruiting asynchrony in contrast to the mainland ; if this were not
,
the case, it is do ub t ful i £ a population of the fig w asp s ( mutualistic seed

predators ) could survive on the small tree population since th e y can live
only a few days outside of the fruit ( 1 78 ) . W he r e toxic compounds in seeds
are only functional in that context, we may exp e ct them to be qu ickly se
lected against on islands lacking seed predators. However, the Hawa iian
endemic Erythrina monosperma is still toxic to eight introduced species of
bruchids ( 33 ) . ( There are no native bruchids on Hawaii, though there are
native seed-eating weevils, 245.) It is interesting to conjecture what a com
petent finch species would do to the trend for larger seeds and less dispersal
observed in Pacific island Com posi tae (44).
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consumption by small mammals. Dominance relations of red and
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Ecology of eastern white pine 4. Adkisson, P. L . , Hanna, R . L . . Bai
seed caches made by small forest ley, C. F. 1964. Estimates of the
mammals. Ecology 5 1 : 27 1-78 numbers of Heliothis larvae per
484 J ANZEN
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Loblolly pine seed production rana ( Lepi doptera: Tortricidae)
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7. Andres, L. A., Angalet, G. W. 1963. 18. B artholom ew, B. 1 9 70. Bare zone
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Entomol. 56:33 3-40 ulations on California grassland.
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the B ruchidae-I. General consid substitution of h ard seeds for
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ecology of the plum curculio, seeds. Natul'e 229; 136-37
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i n th e Niagara Peninsula, Ontario. Ancylostomia stereorea (ZeIt ) ,
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Janzen, D. H. (1971) - Seed Predation by Animals.

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SEED PREDATION BY ANIMALS 4033

PREDISPERSAL SEED PREDATION

seed predator may be set by the density of seed crops, irrespective of

very effective at prey population regulation. This leads one to conclude

POSTDISPERSAL SEED PREDATION

(2 ) , but tropical scatter-hoarding agoutis ( Dasyprocta) may generate

SEED PREDATOR OR DISPERSAL AGENT?

Whether a seed eater is a predator or a dispersal agent depends in great

CHEMICAL ASPECTS OF HOST SPECIFICITY

Fruit clwracteristics ..oC-The primary adaptive functions of fruit morphol­

eralists, they are probably not as important as internal seed chemistry in

Internal seed chemistry .-D irect toxicity may be based on 1 to 10% or

may be reared on partly or completely defined diets ( 105,183, 214, 215 ) , at

Lack of toxicity.-Plants involved i n population and communitywide

predator satiation ( usually temperate-zone trees, but perhaps southeast

Caloric content.-Some vertebrates may harvest seeds largely as a func­

Variance in susceptibility.-Entirely from data on agricultural plants

At the population and community level.-Predat or satiation is undoubt­

edly responsible for the spectacular coevolutionary (somewhat cyclical)

COMPETITION BETWEEN SEED PREDATORS

The density of seed predator populations and communities seems gener­

PARASITES AND PREDATORS OF SEED PREDATORS

The insects preying on large temperate-zone seed crops characteristi­

SEED PREDATORS AND FOREST COMPOSITION

SEED PREDATORS ON ISLANDS

the case, it is do ub t ful i £ a population of the fig w asp s ( mutualistic seed

tera : Curculionidae) and the 12

dae, Co leopte ra) . J. Wash. Acad. Ozarks. J. Forest. 5 3 : 439-41

58. D alke , P . D . 1 9 5 3 . Yields of seeds 72. Elton, C. S. 1924. Periodic fluctua­

foods by gray squirrels. J. Wildl. forage by range rodents. Ecology

geese. J. Wildl. Manage. 30:9-13

1 7 1 . Parnell, J. R 1966. Observations on important shoot. stem. wood.

sile oak <Quercus petraea ) in 1969. Critical factors during the

bean weevil. Ecology 3 4 : 3 0 1-7 truncata Cameron (Hymenop­

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Fruit clwracteristics ..oC-The primary adaptive functions of fruit morphol

Caloric content.-Some vertebrates may harvest seeds largely as a func

At the population and community level.-Predat or satiation is undoubt

The density of seed predator populations and communities seems gener

The insects preying on large temperate-zone seed crops characteristi

58. D alke , P . D . 1 9 5 3 . Yields of seeds 72. Elton, C. S. 1924. Periodic fluctua

bean weevil. Ecology 3 4 : 3 0 1-7 truncata Cameron (Hymenop