Allelopathy by Myrmecophytes: The Ant Azteca as an Allelopathic Agent of Cecropia

Author(s): Daniel H. Janzen

Source: Ecology, Vol. 50, No. 1 (Jan., 1969), pp. 147-153
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1934677 .
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 Winter 1969
LITERATURE CITED
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Skwarra, E. 1934. Okologische Studien uiber Ameisen
und Ameisenpflanzen in Mexiko. R. Leupold, K6-
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Weber, N. A. 1941. The biology of the fungus-growing
REPORTS 147
ants. Part VII. The Barro Colorado Island, Canal
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and their fungus. 1955 Yearbook, Amer. Philosoph.
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tera: Formicidae). Entomol. Soc. Washington, 70:
348-350.

ALLELOPATHY BY MYRMECOPHYTES: THE ANT AZTECA AS AN

ALLELOPATHIC AGENT OF CECROPIA'
DANIEL H. JANZEN
Department of Entomology, The University of Kansas, Lawrence, Kansas 660442
(Accepted for publication August 8, 1968)

Abstract. Azteca ants, that obligatorily live in neotropical second-growth trees of the
genus Cecropia, kill vine ends that begin to climb on the Cecropia trunk. Presumably this
aids the Cecropia tree in its efforts to maintain an emergent position in the general canopy,
and may be one of the selective advantages of being occupied by an Azteca colony. This
system is, in both process and pattern, extremely similar to that of the swollen-thorn acacias
and their ant occupants. In both cases the ants are effectively allelopathic agents, and are
functionally analogous to the chemicals liberated by some plants in their competitive inter-
actions with other plants; like these chemicals, the ants are "produced" at a metabolic cost
to the plant. The ants may, however, be a more efficient allelopathic agent than a chemical
because of their versatility.

Since the original descriptions of the apparently mu-
tualistic association between lowland second-growth trees
in the neotropical genus Cecropia (Moraceae) and the
Azteca ants that obligatorily occupy them (Muller 1874,
1880, Belt 1874), there has been a running argument as
to whether the ants are in any sense "needed" by the
plant (Bequaert 1922, Wheeler 1942, Ihering 1907, Eid-
mann 1944, Bailey 1922, Brown 1960, etc.). That the
argument survives is due to two apparently opposing sets
of observations. Favoring a mutualistic interpretation,
the known species of Cecropia have several apparent
adaptations to the ants: (1) exceptionally large hollow
internodes in which the ants live and tend Homoptera
that produce honeydew (exceptions appear secondarily
derived and are discussed below), (2) regularly occur-
ring thin spots in the internode wall where the worker
and founding queen ants gain entrance to the internode,
and (3) "Mullerian bodies" (lipid, protein and carbo-
1 Contribution No. 1396 of the Department of Ento-
mology, University of Kansas, Lawrence, Kansas. This
study was supported by NSF Grant No. GB-5206 and
the teaching program, of the Organization for Tropical
Studies.
2 Present address: Department of Biology, University
of Chicago, Chicago 60637.
hydrate-rich food bodies) continually growing from highly
modified petiole bases (these tiny bodies are appar-
ently the primary food source of the colony and are
harvested and fed to the larvae). While not all species
of the primarily arboreal, neotropical ant genus A.teca
are involved in this system, at least 10 species are in an
apparently obligate manner (Wheeler 1942, and see also
for a general discussion of the biology of Ateca). The
functional significance of the first two of the three ge-
netically determined morphological traits of Cecropia
listed above has been inadequately "explained" in terms
of structural strengthening (Ihering 1907, Bailey 1922)
and as an accidental by-product of usual developmental
processes (Bailey 1922), respectively. The third trait
cannot be explained in this manner. Unless the ant
colony's presence is considered to be of positive selective
value to the tree, there is no obvious selective basis for
the presence of these three traits.
However, the only immediately obvious aggression by
the ants is against humans that are cutting Cecropia trees,
despite numerous postulations that the ants should pro-
tect the tree from a large variety of herbivores, including
leaf-cutter ants. Further, it is commonplace to encounter
occupied Cecropia with foliage heavily eaten by insects,
and healthy unoccupied Cecropia are apparently not rare

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148
in some habitats. The obvious field experiments needed
to clarify the situation had not been attempted.
This apparently contradictory situation, accompanied
by similar arguments, was the state of the art for all
apparent mutualistic associations between plants with
ants living in them (myrmecophytes), until it was experi-
mentally demonstrated that two Central American swol-
len-thorn acacias (Acacia cornigera and A. collinsii;
Leguminosae) are obligatorily dependent in nature upon
their ant occupants (Pseudomnyrmex spp.; Pseudomyr-
mecinae) for survival to reproductive maturity (Janzen
1967a and unpublished). Further, the ant-acacia myrme-
cophytic relationship is a very highly coevolved one
(Janzen 1966) and applies to all 10 of the normally
occupied swollen-thorn acacia species in Central America.
The demonstration of mutualism between Pscudo11nyr-
wnex and Acacia throws a new light on the Cecropia and
Asteca interaction, especially since the general extent of
herbivore damage to swollen-thorn acacias is very simi-
lar to that of Cecropia. Further, the only immediately
obvious aggression demonstrated by Pseudomyrinex is
against large vertebrates, and like Cecropia, apparently
healthy but unoccupied swollen-thorn acacias are occa-
sionally encountered in nature. However, detailed analy-
sis has shown that Pseudoinyrnmex workers remove most
phytophagous insects and kill the intrusive branches of
vines and shrubs in the acacia canopy. If the ants are
experimentally removed, the combination of greatly in-
creased insect damage and shading from adjacent plants
in the rapidly growing adjacent second growth -severely
stunts the plant and eventually results in death. Further,
a partial "fire-break" may on occasion result from the
ants' pruning activities (Janzen 1967b). Alone, either
browsing by insects or shading will kill the acacia, but
they are faster and more effective when operating to-
gether.
The analogous experiments with Cecropia (and two
other myrmecophytes, Cordia and Triplaris) are presently
in progress in Costa Rica, but I would like to draw at-
tention at this time to one activity of worker Axteca ants
that very likely influences the survival and general health
of the Cecropia they occupy.
By chewing on them, worker A teca kill the shoot tips
_ f .
',.....,
.... A...,.A..... . A.....
i ...........................
ir..'.,.......p
w..... ;...I;,....I
FIG. 1. On the right are two vine ends of a convolvu-
laceous vine that have been chewed off and killed by growing woody emergent, and having large horizontal
Azteca worker ants where the vine touched the Cecropia.
The vine on the right has also had one leaf completely
pruned off and the other severely trimmed. The two vines
on the left and extending up over the top of the photo-
graph are intact examples from the same plant (8 March
1968; Osa Peninsula, Costa Rica).
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REPORTS Ecology, Vol. 50, No. 1
and tendrils of vines that attempt to climb Cecropia
trunks or enter the canopy laterally (Fig. 1); they in
effect make the occupied Cecropia tree capable of alle-
lopathy. Studies of allelopathy (as defined by Molisch
1937) have focused on chemical inhibition of neighboring
plants (Muller, Muller, and Baines 1964, Muller 1966,
Borner 1960, Woods 1960) and the mechanics of the
process. However, the phenomenon may be generalized
as a competitive process (of an interference type), or
model, whereby one organism is responsible for the input
into the environment of an object that is inimical to the
development of a competitor (i.e., competitive interfer-
ence where the adaptive value is in the elimination of a
competitor). Here, the severe destruction of vegetation
around swollen-thorn acacias by obligate acacia ants (Jan-
zen 1967a) is clearly analogous to the inhibition of herba-
ceous growth around the shrubs Arteniesia and Salvia
by volatile compounds (Muller et al. 1964, Muller 1966)
and other classical cases of allelopathy. As is likely the
case with the chemicals active in allelopathy, the ants are
"produced" by the plant in a very real sense, in respect
to the cost of maintaining them and the selection for
traits promoting their presence.
Barnwell (1967), while discussing activity patterns,
first described damage to other plants by Azteca cf.
alfari on Cecropia obtusifolia (near Puerto Viejo, Heredia
Prov., Costa Rica): "the ants . . . were cutting deeply
into the stem and leaves of a cucurbit . . . where it
had contacted the Cecropia. Similarly, at the C. peltata
in Guanacaste the leaves of an adjacent shrub had been
neatly cut back from the lower trunk of the tree." He
then postulated that "there is the possibility that the
cutting back of overgrowth contributes to the survival
and reproduction of Cecropia . . ." I would like to sup-
port this hypothesis as a special case of allelopathic
interference competition, and discuss its relationship to
the hitherto unsatisfactorily explained aggression to
plants shown by several genera of ants occupying myrme-
cophytes other than Cecropia and Acacia.
METHODS
The speed, effectiveness and specificity of Azteca cf.
alfari workers in removing vines from Cecropia peltata
and C. sandersonianac was examined 4-6 miles SE of
Rincon, Osa Peninsula, Puntarenas Prov., Costa Rica.
Forty vine branches 1-3 m long were carefully untwined
from shrubs within 1 m of the Cecropia trunk, and lightly
twined around the trunks of 18 occupied Cecropia (the
remainder of the vine plant was undisturbed). This is
directly analogous to placing plants within the range of
the allelopathic compounds liberated by some plants.
That this is a reasonable experiment is shown by the
= .S.C::,',.Aj~~~~~~~~~~~~~~f,:.
presence of many chewed and blackened vine ends ad-
jacent to the lower trunk of occupied Cecropia. As is
the case with swollen-thorn acacias (Janzen 1967a),
some trunks are literally ringed by dead ends of vine
shoot tips and tendrils and project out of a solid mat of
vines. These dead vine ends may be 1-20 cm from
the Cecropia trunk as a result of stem movements after
being chewed off by the ants. Further, vines are fre-
quently present on unoccupied Cecropia and generally
absent from occupied Cecropia. Being a strong and fast-
petioles and branches, it makes an excellent vine standard
and quickly becomes one when it loses its ant colony
through natural means (Fig. 2, upper photograph).
The fate of these 40 vines was observed from 1 to 8
March 1968 (Table 1). This period is about 2 months
after the beginning of the dry season (see similar rainfall
Winter 1969 REPORTS 149

* :~~~~~~~~~~~~~~~~~~~~~~~~~~....
.I..... ......
. .................... ........ ..... .:..i... ... ..:..;
.....

Aersnaiemmbronioae uncupe pouato of Ceroi (udsrbd at 20 0melvto

hinCsaRc(2 Ma198. Temsssadlcesothbrneswudot liel be prsn if th tre
wer ocupe (see text).

FIG. 2. Upper: On the left is a 6 m unoccupied Ce cropia covered with vines, as is commonly the case with
unoccupied Cecropia. On the right is a Cecropia heavily occupied by Aztcca. Both trees are approximately 5
years old and are growing in a marsh near Dominical, Puntarenas Prov., Costa Rica (25 May 1968). Lower:

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150 REPORTS Ecology, Vol. 50, No. I
TABLE 1. Fate of intact shoot tips and tendrils on the by any one of several species of obligate acacia ant.
ends of 40 vine branches artificially twined around 18 Pseudomyrmnex have larger mandibles, are generally
Cecropia peltata and C. sandersoniana trunks 1-2 m heavier in build (about 6 mm long as compared with
above the ground (1 March 1968)a 2-4 mm), and often sting the vine (perhaps causing some
necrosis). There is no evidence that defensive com-
pounds are placed in the plant wound by the Azteca
Condition 3 March 8 March
workers, but this is possible since they do this when
Not yet damaged............. . 25% 10% biting humans. If they do, then the process becomes one
Being chewed on at the time of recording. . 20% 0% almost identical to allelopathy in the classical sense.
Chewed on............................ 23% 8% Failures of the ants to kill a vine tendril or shoot tip
Dead by ant chewing.................. 25% 75% in the 8-day period were associated with several vari-
Blown off.......... ............. 8% 8% ables. (It is doubtful if any would have remained alive
indefinitely since other older vine ends within a few
aThe Cecropia were 4-12 m tall and all occupied to some degree by an active centimeters of the trunk had been killed by the ants).
colony of Azteca cf. alfari. Only those vine ends with some part of the terminal The size of the ant colony was positively correlated with
20 cm in contact with the trunk were recorded.
the shoot tip mortality rate. The larger the colony, the
heavier the nocturnal activity on the surface of the stem;
records from Golfito; Janzen 1967c), but vine growth most of the chewing appeared to occur at night (again,
was still active since it generally rains at least once a this may be a peculiarity of the dry season since the
week at this site. The sample size was slightly reduced general activity of the colony is much lower in the dry
by the wind blowing some vines off, since no string was season than the wet). The higher the Cecropia canopy
used to tie them down and the normal shape of the vine above the contact point of the vine, the slower the ants
was not distorted to make it hold onto the tree. The were to encounter it (the primary ant activity is in the
species of vines were chosen purely by their adjacence canopy) ; it may be noted, however, that the more emer-
to the Cecropia and ease of removal from their support gent the Cecropia canopy, the less danger there is of
without damage. At least 10 families were represented vines overtaking and completely covering the canopy
(Passifloraceae, Cucurbitaceae, Bigoniaceae, Convolvula- (the same is true of swollen-thorn acacias). In effect,
ceae, Fabaceae, Caesalpinaceae, Apocynaceae, Composi- there is a lessening of allelopathic activity with increase
tae, Asclepiadaceae, Malpighiaceae, etc.) and at least in vertical distance of the Cecropia canopy from other
18 species. They ranged from very tough and hirsute plants in the community. Observation of "natural experi-
(e.g., Asclepiadaceae) to glabrous and weak (e.g., Con- ments" similar to those done in this study indicate that
volvulaceae). No controls (in the form of vines twined vine tendrils or shoot tips are always killed when they
around other trees) were used because the damaging enter the canopy of a heavily occupied Cecropia no matter
agent was under observation and because on many pre- how tall it is. It should be emphasized that as with
vious occasions, vines have been manipulated in this man- Pseudomyrmex in swollen-thorn acacias, there appears
ner with no mortality of their shoot tips. to be individual colony variation in aggressiveness that is
unrelated to the above factors, or to colony age, vegeta-
RESULTS tion type, time of year, etc. In short, there is moderate
The shoot tip mortality recorded in Table 1 shows variation in allelopathic abilities of occupied Cecropia,
clearly that the chewing by ASzteca workers can rapidly and part of this is apparently beyond the control of the
kill shoot tips (apices) and tendrils; after 8 days, 75%o plant.
were dead by chewing, 8% had been chewed on but were All the vine species that had not yet been killed were
still green, and 10%o were contacting the tree but not represented among those that had been killed. Very
yet damaged. On 13 of the 18 trees the ants were ob- hirsute vines were first stripped of their hairs at the
served chewing on the vine ends, and on the others, it point of contact with the trunk and then gnawed off.
was obvious by mandible marks and the characteristic When latex was produced (e.g., Asclepiadaceae, Apocyna-
blackening and shriveling of the vine stem that the dam- ceae), it dried and the ants chewed it off. Where vines
age had been done by the ants and not some other insect. were about equally attacked by ants in respect to num-
The blackening occurred only where the vine had been bers and timing, it was clear that the fragile ends of
chewed on, not necessarily at its points of contact with Bignoniaceae and Convolvulacea were killed two to five
the Cecropia trunk. The ants lined up in parallel at times as fast as the tough ends of Compositae and Mal-
right angles to the vine shoot tip where it contacted the pighiaceae. However, the species in the former two
Cecropia trunk and generally chewed on it until it broke families grow much faster than do those in the latter
off, or the contacting portion was removed or wilted and it is not clear which, if either, would be more suc-
away. When the vine contacted the Cecro pia trunk cessful in climbing on the Cecropia when confronted by
10-20 cm below the tip, the ants were slower to move a weak ant colony.
out onto the stem and kill the tip, but sometimes did. There was no evidence that a toxic compound in the
More often, they chewed through the stem where it con- Cecropia bark was killing the vines, though this would
tacted the trunk, even though the stem was much tougher be a reasonable defense against vines. In that A-zteca
there than near the tip. only kill vines when very close to the trunk, rather than
Chewing was discontinuous; no ants were observed at distances up to 2 m as does Pseitdomyrmex on swollen-
active at the vines during the midday hours (this may, thorn acacias, they are more analogous to an allelopathic
however, have been influenced by the dry season when compond that is only in the bark (and thus effective only
midday activity is reduced). Workers of all sizes chewed against vines and epiphytes) than one that is liberated
on the vines. Azteca workers are much slower than widely through leaching or volatilization.
Pseudomnyrmex ferruginea (Janzen 1967a) in re-noving
vines of equal sizes. A vine that would require about 4& DISCUSSION
hours to be removed by an active Aztecta colony would Throughout the second growth in lowland Central
be removed from a swollen-thorn acacia in about 8 hours America the population of Cecropia, like that of swollen-

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Winter 1969
thorn acacias (Janzen 1967a), is made up of two sub-
populations. There are the numerous 0.2 to 1.5-m-tall
unoccupied or very weakly occupied seedlings or sucker
shoots, and the maturing or reproductive individuals that
are usually, though not invariably, occupied by a large
Azteca colony. Again as with swollen-thorn acacias,
most of these small trees are not noticed by casual ob-
servation because they are covered by vines or heavily
obscured by dense shrubs. It appears to be a rare event
when one of these stunted plants "breaks free" from the
shade of the adjacent or covering plants and matures.
This does not, of course, apply equally to all types of
vegetation. On open gravel bars in rivers, there are
few other plants and the young unoccupied Cecropia
often grow unhindered by other plants. However, this
is not the case on more permanent river edges where there
is intense competition for light between Cecropia trees
and both upright plants and vines.
To the degree that there are vines in the surrounding
plant community, there is a definite opportunity for Az-
teca to influence Cecropia survival and reproduction
positively. This observation is based on the heavy dam-
age to vines contacting recently occupied small Cecropia,
the damage to vines reported in this paper, the definite
negative effect that shading has on Cecropia growth rates
and reproduction (when heavily shaded, they do not
reach reproductive maturity), the general absence of
vines from heavily occupied Cecr-opia, and their general
presence on unoccupied Cecropia. If this positive in-
fluence occurs, then it can provide the selective pressure
for all the traits of Cecropia that maximize ant colony
development: Miillerian bodies for food, regular thin
spots in the internode wall that facilitate founding queen
entrance and colony establishment, 'Largeinternal cavities
(domatia), no biochemical defense against the honeydew-
producing Homoptera, leaf retention during the dry sea-
son. These traits yield the same end product as would
an allelopathic chemical in the tree bark, with the addi-
tional advantage that the ants may also deter some her-
bivorous insects as is clearly the case with obligate
acacia-ants.
While the ants' reaction to epiphytes was not examined,
it is suspected that they would remove them in the seed-
ling stage since angiosperm, fern and lichen epiphytes
are characteristically absent from heavily occupied Ce-
cropia of all ages. The fact, however, that Cecro-
pia grow rapidly (up to 4 m/year in areas with a
short dry season), and die after 10-15 years, may reduce
the possibility of epiphyte development; other second-
growth trees also often lack epiphytes until they have
grown to the point where branch shedding is reduced.
On the other hand, in dense mixed stands where Cecropia
makes up only a small percentage of the biomass, and
the regeneration is over 10 years old, virtually all trees
over 10 m tall have epiphytes and/or vines on them ex-
cept Cecropia. Cecropia is often the only tree species
that can consistently be easily cut and pulled free from
the general vegetation.
Epiphytes appear to be abundant on Cecropia only at
elevations too high for the ant. In central Costa Rica
(16-23 km south of Cartago; 1900-2000 m elev.) a
small population of an undescribed species of Cecropia
is apparently at too high an elevation for Azteca colonies
to survive. This species has all the traits of a normal
Cecropia and is clearly an isolated segregate from a
lowland species. The trees are festooned with epiphytes
of many types (Fig. 2, lower). That they are not
covered with vines is correlated with the general absence
of high-climbing vine species at this elevation and the
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REPORTS 151
much slower growth rate, and lower density of the vines
that are there, as compared with the lowlands.
It is not uncommon to find a large and occupied Ce-
cropia with a moderate number of large (old) vine stems
in it. As is the case with swollen-thorn acacias, once a
large vine has become established, it is difficult for the
ants to remove it. Psezdomiyruiiex eventually kills almost
any vine, but this is primarily due to their greater ag-
gressiveness than Azieca. On the other hand, old estab-
lished vines on occupied Cecropia show little new growth;
when a new tendril or shoot tip comes in contact with
the Cecropia leaf or trunk, the ants kill it. Maturing
Cecropia may become standards for vines through tem-
porary unoccupation (colony abandonment during the
dry season, colony death with later recolonization) or by
being occupied by a weak ant colony or even a strong
one that for unexplained reasons is not aggressive to
vines. Primarily due to growing in a habitat generally
lacking vines, unoccupied maturing Cecropia on occasion
lack vines.
Azteca are much less effective than Pseudonzyrmiex in
removing foliage from branches of the neighboring erect
vegetation because they usually chew the vegetation only
at its point of contact with the Cecropia. It is common
to find a leaf neatly trimmed off along its margin of con-
tact with the Cecropia trunk. However, maturing Ce-
cropia are often very emergent during the first half of
their life (in contrast with swollen-thorn acacias which
are usually canopy members or no more than 1-2 m emer-
gent). They therefore rarely have foreign branches in
contact with their canopy. On the other hand, small and
unoccupied, or only recently occupied, Cecropia are often
thoroughly immersed in the dense general canopy of 1 to
3-year-old regeneration, bringing many of the "offending"
branches directly in contact with the Cecropia canopy
where the ants then do work on them.
Since Azteca do not extend their pruning activities to
a small circular area under the Cecropia (as does Pseudo-
myrrnex), it is doubtful if this activity is functional in
reducing fire damage. However, Cecropia usually live
in sufficiently wet sites such that fire is quite rare in
contrast to Acacia habitats (perhaps adding support to
the hypothesis that the "fire-break" function of the basal
circle is more than purely accidental; c.f. Janzen 1967b).
It may, however, be noted that allelopathy in the classi-
cal sense, where it results in a clear area around a woody
plant (Muller et al. 1964), may have a "fire-break"
function.
I have observed the allelopathic habit in at least five
species of Azteca occupying at least five species of Ce-
cropia from Panama to coastal central Mexico. Over
this wide range some Cecropia occur in habitats where
vines are very rare; it is therefore difficult to observe
vine damage, and further, in these habitats allelopathy
by the ants is probably not a primary function of the
ant colony. In other habitats, primarily the wet and
well-drained sites, vines are a very maj or competitor
with woody plants and thus allelopathic activity is likely
of major importance.
The origin of the plant-chewing habit in Pseudoinyrniex
appears to be simply an extension of the general aggres-
sion toward any foreign object (Janzen 1967a, b), and
there is no reason to believe otherwise for Azteca (de-
spite comments to the contrary, Bequaert 1922, Barnwell
1967). As with Pseudomyrmex there is no evidence
that it is related to food gathering since it is least in-
tensively pursued when both food and moisture production
by the myrmecophyte is lowest (dry season). The ants
stop as soon as the vine or branch no longer touches the
152 REPORTS
Cecropia, and the ants have never been observed to "raid"
the surrounding vegetation only a few centimeters from
the base of the trunk. Ants frequently fall from the
canopy onto the vegetation below and then return to the
trunk of the Cecropia without chewing on the vegetation.
Finally, an AS-teca colony deprived of its usual food
sources (by cutting the tree) does not forage on the
vegetation the cut trunk is lying on. These observations
do not preclude the possibility that they drink the plant
fluids at the time of chewing on the vine.
It may be noted that this system solves the difficulty
in allelopathy of the evolution of chemicals that are toxic
to other plants but not toxic to the producer. The ants
themselves are not toxic to the myrmecophytes, but this
does require a behavioral adapation on the part of the
ant. In the swollen-thorn acacias this is imperfectly ( ?)
developed in that obligate acacia-ants kill seedling swollen-
thorn acacias under the parent, but not root sucker shoots
(which have, however, nearly adult foliage). In an
exceptional case, however, Pseudoiinirniicx fe1rrtolyea, in
the drier portions of Nicaragua, even prune the shoot
tips of the Acacia coll/iisii they occupy, producing a very
shrubby plant.
The allelopathic habit of ants obligate to a myrmeco-
phyte is one of the clearer examples of convergence by
the ants in the coevolution of ant and higher plant inter-
actions. This makes it even more likely that it is func-
tional for the plant since ants in general do not attack
vegetation. In ad(lditioll to Pseodoiiiyriniex in swollen-
thorn acacias and Aziteca in Cecropia, the distantly re-
lated Azteca. species in the Central American late second-
growth tree Cordia alliodora (Boraginaceae) also vigor-
ously attack vine ends in the canopy. Distantly related
I'SCodo1yr1nre species occupying the Central and South
American Triplaris (a river-edlge tree in the Polygona-
ceae) kill vines entering the canopy and clear the green
vegetation from below the tree in exactly the same man-
ner as do the obligate acacia-ants in Acacia (Ule 1907;
the trees are called "Palo Santo," apparently referring
to their extremely clean appearance; J. Idrobo, pers.
comm.). There is apparently some confusion on this
since Wheeler and Darlington (1930) state emphatically
that the ants do not clear an area under Triplaris. Other
sidel0tv'me- il Tachigalia (a South American river-
side and old second-growth tree in the Leguminosae) do
the same thing, and it is very clear that the plant must
free itself from the shade of neighboring plants for both
ant colony and tree survival (Wheeler 1921). In the
Belgian Congo, a sister genus to Pseudoitzyroiiex, Pachy-
sinoa, has a very similar relationship with Barteria (a
weedy second-growth tree in the Flacourtiaceae or Passi-
floraceae), as Pseotdoinyrnex does with Acacia. Here,
also, the ants kill the vegetation growing under the tree
(Bequaert 1922, Kohl 1909), and the pure stands of
Barteria in old elephant feeding grounds (Jones 1955)
probably are the result of this (as with the pure stands
of swollen-thorn acacia in Central America, janzen 1967a,
b). Bequaert (1922) saw the function of the vegetation
clearing by Pach ysima as allowing the ants to find prey
insects "so much easier and quicker;" the problem with
this interpretation is that Pachysimlia, like Pseudomyr-
wnex.,subsists on food bodies from Barteria, not prey in-
sects. Mfacaranga (Euphorbiaceae), the southeast Asian
highly convergent analog of Cecropia, "never" has epi-
phytes on it (Ridley 1910), and this is very likely due to
aggressiveness to plants by their occupant ant Cremato-
gaster (Myrmicinae).
Destruction of vegetation around ant nests in general
is obvious in two other cases. The grassland nests of the
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Ecology, Vol. 50, No. I
harvester ant, Pogonoiniyric.r occiden talis, are centered
in a 1- to 3-m-diameter bare circle inl dense grass. While
this bare circle may be purely the result of seed-gathering
activity by these ants, it is also heavily reinforced by the
ants chewing down the growing vegetation. The func-
tional significance of this may be in creating a small
"fire-break" around the entrance (Cole 1932), as well
as increasing insolation of the ant mound itself. Secondly,
the mound-building forest ant, Formnica exsectoides, in
the northeastern United States has been shown to prune
or kill by mauling (accompanied at times by injection of
fluids from the gaster) the small trees and herbaceous
vegetation to distances of several meters from the nest
(Andrews 1928, Peirson 1922). The resultant circular
clear area is very likely functional in increasing insolation
of the nest mound, especially during the spring and fall
when such insolation could substantially increase the tem-
perature of the nest. While a living plant's competition
with another plant is not involved here, the result is very
similar to more conventional allelopathic systems, and
stresses the similarity between an ant colony and a plant
in their functional interactions with the environment.
CONCLUSION
In summary, there is clearly at least one way that the
Az/eca ants that occupy Gecropia can positively influence
the tree. This damage to vines, and at times leaves front
adjacent erect plants, falls clearly into a generalized model
of the special case of interference competition known
as allelopathy. As with the chemicals produced by a
plant in the allelopathy, the maintenance of an ant colony
is a definite drain on the energy budget of the tree. Fur-
ther, the ants can be seen as analogous to the more comi-
mon defense mechanisms of plants in their interactions
with other organisms. This system differs from the
usual cases of allelopathy in that certain components of
the system's effectiveness are probably beyond the "selec-
tive reach" of the plant. However, the system may he
more complicated than appears on the surface; for ex-
ample, it is quite possible that the food bodies of the
Cecropia contain compounds that influence the general
aggressiveness or irritability of the ants. The ants differ
from the usual chemicals associated with allelopathy in
that it may be more difficult to evolve a specific defense
against them than against a specific chemical, and they
are in addition a very efficient insecticide (as least in the
case of swollen-thorn acacias, and likely in other myrm-ne-
cophytes to a lesser degree).
LITERATURE CITED
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building ants. Amer. Naturalist 62: 63-75.
Bailey, I. W. 1922. Notes on neotropical ant-plants.
I. Cecropia angulata, sp. nov. Bot. Gaz. 74: 369-391.
Barnwell, F. H. 1967. Daily patterns in the activity
of the arboreal ant, Azteca alfari. Ecology 48: 991-
993.
Belt, T. 1874. The naturalist in Nicaragua. Bumpas,
London. 403 p.
Bequaert, J. 1922. Ants in their diverse relations to
the plant world. In Ants of the American Museum
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cology of Africa. Bull. Amer. Mus. Natur. Hist. 45:
1-1139.
Burner, H. 1960. Liberation of organic substances
from higher plants and their role in the soil sickness
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mex occidentalis Cr., to its habitat. Ohio J. Sci. 32:
133-146.
Eidmann, H. 1944. Zur kenntnis der Okologie von
Azteca muelleri Em. (Hym. Formicidae) ein Beitrage
a urn problem der Myrmecophyten. Zool. Jahr. Abt.
Syst. Okol. Geogr. Tier. 77: 1-48.
Ihering, H. v. 1907. Die Cecropien und ihre Schutza-
meisen. Engler's Bot. Jahrb. 39: 666-714.
Janzen, D. H. 1966. Coevolution of mutualism be-
tween ants and acacias in Central America. Evolu-
tion 20: 249-275.
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(Acacia cornigera L.) with an ant inhabitant (Pseudo-
rnyrmnexferruginea F. Smith) in eastern Mexico. Univ.
Kansas Sci. Bull. 47: 315-558.
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Jones, E. W. 1955. Ecological studies on the rain
forest of southern Nigeria. IV. The plateau forest
of the Okomu Forest Reserve. J. Ecol. 43: 564-594.
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Afrika mit besonderer Berucksichtigung ihrer bio-
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REPORTS 153
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DIVERSITY IN TERRESTRIAL CAVE COMMUNITIES

THOMAS L. POULSON AND DAVID C. CULVER1
Yale University and Cave Research Foundationl
(Received April 14, 1968. Accepted for publication September 18, 1968)

Abstract. Local species diversity of terrestrial arthropods was determined from a com-
bination of trapping and census in an area of variable passage type in Flint Ridge Cave
System in Mammoth Cave National Park, Kentucky. We measured evaporative rate, sub-
strate moisture, substrate organic content, predictability and stability of food and microclimate,
substrate diversity, and intensity of flooding. We found significant correlations of species
diversity with substrate diversity, substrate organic content, and intensity of flooding.

Biologists have known for many years that there are
large differences in the number of species and their rela-
tive abundances on both a local and a geographic scale.
However, it was not until the development of informa-
tion theory (Shannon 1949) that biologists had at their
disposal a measure of diversity that took into account
both the numbers of species and their relative abundances
and was free of any assumptions concerning the theo-
retical distribution of such an index. MacArthur (1965)
recognized two components of diversity, within-habitat
and between-habitat diversity, and pointed out that within-
habitat diversity must be understood before between.
habitat diversity can be considered. Field studies on the
causes of differences in within-habitat diversity are scanty
and primarily deal with birds (MacArthur and Mac-
1 This study was supported by United States Public
Health Service grant GM 12,231 to T. L. P. We would
like to thank the National Park Service for the use of
various facilities. Stewart Peck offered many helpful
suggestions.
Arthur 1961) and lizards (Pianka, 1967). Some per-
tinent studies have dealt with invertebrate groups such
as fresh-water cladocerans (Goulden 1966, Whiteside
and Harmsworth 1967) and tree insects (Southwood
1961).
The terrestrial cave community is readily amenable to
an analysis of within-habitat diversity. Since many en-
vironmental factors are constant and the fauna is rather
depauperate, it is possible to consider the entire macro-
scopic community. It is our intent to examine a terres-
trial cave community consisting of various arthropods in
order to determine whether any generalities are possible
at the community level and to ascertain the relationship
of our results with those of previous workers who have
considered other groups in different habitats.
METHODS ANI) MATERIALS
Field work was done in July of 1966 and August of
1967 in Mammoth Cave National Park, a part of the
Central Kentucky Karst on the west flank of the CGin-

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