Nagano, T. F. & Burgos, M. G.

Bio 165 – Group 4

Experimental Zoogeography of Islands

D.S. Simberloff & E. O. Wilson

Part I: Defaunation and Monitoring Techniques

The experiment tests MacArthur and Wilson’s (1963,1967) quantitative theory of island biogeography
that postulates that an equilibrium number of species on an island determined by the intersection of
immigration and extinction curves drawn as a function of the number of species already present.

The Experimental Islands (Florida Keys)

Nagano, T. F. & Burgos, M. G.
Bio 165 – Group 4

The islands with diameter 10-20 m are 5-10 m tall and usually consist entirely of one to several
Rhizophora mangle tress, with rarely a small black mangrove bush (Avicennia nitida). Occasionally,
in eareas of weak tide, the trees are surrounded by small areas of supratidal mud and sand.
The experimental islands must adhere to the following conditions:
1. There must be enough of the islands for replication and variation in distance to the nearest source
area.
2. The animal diversity must be sufficient to allow statistical treatment
3. The extremely small size of the islands compensates for their relative nearness to source areas and
therefore produces a distance effect.

Defaunation
Fumigants highly soluble in water could not be used. Hence, the following four relatively insoluble
fumigants were tried in field tests with R. mangle trees: methyl bromide, 34% acrylonitrile + 66%
carbon tetrachloride, ethylene oxide, and sulfuryl fluoride. Ethylene oxide, sulfuryl fluoride, and 34%
acrylonitrile + 66% carbon tetrachloride caused extensive irreversible damage to R. mangle and were
rejected. However, methyl bromide at 22 kg/1000 m3 for 2 hr caused browning of only about 10% of
the leaves 1 week after treatment, with no visible later effects. This duration and concentration killed
all the mangrove inhabitants (including resistant forms such as roach eggs and lepidopteran pupae).
Two islands (E6 and E10) were selected to serve as control islands; and they were censused at the
same time as the experimental islands.
Nagano, T. F. & Burgos, M. G.
Bio 165 – Group 4

A cuboidal scaffolding framework was used to

construct a tent over the experimental islands. Methyl
bromide was introduces from 45 kg cylinders
through the tent wall at opposite sides of the island,
2.5 m above the water, and into a metal tub mounted
with an electric fan to disperse the gas. The methyl
bromide concentration was gradually built up from 0
to 22 kg/ 1000 m3 over 30 min, then kept at 22-25 kg/
1000 m3 for 2 hr. All islands were examined
immediately after tent removal. No living animals at
all were found on E1, E2, E8, and E9. A single live
curculionid larva was found on E3. On ST2, 1 living
and at least 100 dead polyxenid millipedes were
found.

It was discovered that the rate of discovery of new species during each censusing period declined
approximately linearly to near 0 at about 14 man-hours of searching. Because of the simplicity and
very small size of the experimental islands, and because several nocturnal censuses located no new
species, it was assumed that all colonist having crepuscular or nocturnal activity were being recorded
during the diurnal monitoring.

Part II: Colonization of Empty Islands

o Propagule – the minimum number of individuals of a species capable of breeding and
population increase under ideal conditions (i.e. unlimited food supply and proper habitat, no
predators, etc.)
o Colonization – the existence of at least one propagule of a species on an island
o Extinction – the disappearance of a species from an island
o Invasion – the arrival of one or more propagules on a n island
o Immigration – the arrival of a propagule on an island unoccupied by the species (in units of
species/time). Every immigrant is a colonist.
It was assumed that an adult female, an adult of indeterminate sex, and an immature animal each imply
the presence of a propagule. Adult males are not considered colonists. In addition, all Diptera were
excluded because monitoring of flies proved too difficult to warrant faith in the accuracy of species
counts.
Nagano, T. F. & Burgos, M. G.
Bio 165 – Group 4

Seasonality
The Florida Keys are subtropical, with the mean temperature of the coldest month (20.9 degrees
Celsius) only 7.7 degrees Celsius lower than that of the warmest month. In addition, frost has never
been recorded and wind is relatively constant over the year in both speed and direction. Hence, most
mangrove arthropod species were found active throughout the year and all life stages were present
during every month.

Patterns of Colonization
Because the experimental system involves only one plant, succession (a progression of discrete and
relatively stable communities) did not occur. This does not imply lack of order in colonization; the
invasion of species was surprisingly regular. But, it was not accompanied by wholesale extinction of
distinct animal associations.
The earliest immigrants on all islands included both strong fliers (moths and wasps) and weak fliers
or nonfliers (psocopterans, chrysopids, and spiders), the latter built up large populations more
rapidly and became numerically dominant. Many of the early invaders was transported via wind
dispersal.
==========================Variable Colonization Patterns===========================
Psocopterans
Colonization of the early weak fliers was more variably than the other classes of arthropods. A total
of 24 psocopteran species were involved; Psocidus and Peripsocus were generally more prominent
than other psocopterans. Regardless, there was little correlation among psocopterans found on
different island. Many psocopterans persisted for less than a month. Ants, such as Pseudomyrmex
elongatus and P. flavidula, and running and jumping spiders, can eat psocopterans.
Spiders
The majority of the 36 spidder species observed followed the pattern described for psocopterans –
they immigrated readily and became extinct quickly. However, Eustala, Tetragnatha, Leucauge
venusta, Hentzia palmarum, and Aysha velox colonized most of the islands and persisted for at least
several months.
Wasps
Some wasps appeared on one or two islands and vanished rapidly. While a few, notably Pachodynerus
nasidens, Sclerodoma macrogaster, and Calliephialtes ferrugineus colonized many islands and
persisted for long intervals.
Mites
Mites did not invaded as early as wasps but displayed the same pattern of many short-lived species.
Most Acarina species disappeared within a month, while Amblyseius and Galumna were omnipresent
and populations long-lived.
Nagano, T. F. & Burgos, M. G.
Bio 165 – Group 4

===========================Regular Colonization Patterns==========================

Thrips
Only 5 thrip species colonized, 4 of which were widespread. Almost all invasions occurred 4-5
months after defaunation; there was no consistent order of colonization – all species commonly
immigrated at about the same time. Most thrip colonizations endured for at least 3 months; large
populations rarely persisted as the late colonizing ants may have attacked thrips.
Lepidopterans
Eight lepidopterans colonized the experimental islands; all were recorded more than once and six
were widespread.
Order of invasion:
Phocides batabano + Bema ?yyda + Ecdytolopha -> + Nemapogon -> + Alarodia slossoniae + Automeris
Orthopterans
Nine invaded experimental islands; if two very near islands (E2 and E7) are discounted, only four
species were involved and all appear capable of early invasion, have multiple records and were rarely
extinguished. Cyrtoxipha confusa (cricket) and Latiblattella (roach) produced large populations.
Ants
They displayed the most orderly pattern of colonization and are the numerically and energetically
dominant animals on all small mangrove islands. Ants colonized later than most other taxa.
Equal island size, from distant islands to near shore:
Crematogaster ashmeadi -> + P. elongatus -> + Paracryptocerus varians + Tapinoma littorale +
Camponotus -> + P. flavidula <or/+> Monomorium floricola <or/+> Xenomyrmex floridanus
Equal distance from source area, islands of increasing size:
C. ashmeadi -> + P. elongatus -> + Paracryptocerus + Tapinoma + Camponotus + Xenomyrmex +
Monomorium
CONCLUSION: The ability to colonize increasingly smaller islands parallels closely the ability to
colonize increasingly distant islands.
Only 3 extinctions were observed, and 2 were of species unable to nest in R. mangle.

Colonization Curves
The number of species on the control islands (before and after defaunation of the experimental
islands) did not change significantly; however, the species composition varied considerably, implying
that the number of species, S, approaches a dynamic equilibrium value, 𝑆̌.
Nagano, T. F. & Burgos, M. G.
Bio 165 – Group 4

Three evidences for the existence of dynamic equilibrium value, 𝑆̌:

1. S on the control islands did not change greatly from the beginning to the end of the year-long
experimental period.

2. Untreated islands with similar area and distance from source have similar 𝑆̌ (note E3 and ST2
before defaunation.
3. Increase of species present in all islands to approximately the same number as before defaunation,
and then rough oscillation about this number.
SPECIAL CASES:
E1
 The island was so distant from its source area that a few early invaders were able to build up
large populations before competitors arrived. Interactions became important before even a
small fraction of the non-interaction 𝑆̌ was achieved. As such, E1 colonization curve will
ascend slowly and irregularly to an equilibrium near pre-defaunation 𝑆̌.
 Non-interaction 𝑆̌ should decrease as the distance of the island from the source increase
because of decreases in invasion rate. The time necessary to reach any percentage of 𝑆̌ should
increase with increasing distance from source. If this time is sufficiently long, the few early
colonists are able to produce large enough populations to interact with later immigrants.
E7
 85% of the tree was killed by fumigation. As such, the relative proportions of microhabitats
changed drastically until there was more dead wood and bark and less leafy canopy. The
expected number of species would therefore change because different microhabitats support
different number of species. Dead bark shelters numerous species while mangrove leaves
Nagano, T. F. & Burgos, M. G.
Bio 165 – Group 4

rarely support more than four species on a single island; as such, E7 colonization curve rose
to a higher percentage than the original 𝑆̌ (135%). Arachnids and psocopterans (both bark
dwellers) were largely responsible for the higher S.

Immigration and Extinction Rates

The observed immigration and extinction curves for all islands were highly variable. The turnover
rates are surprisingly high, in the vicinity of 1% of the equilibrial species number per day or higher.
This is consistent with the MacArthur-Wilson model, which predicts that the turnover rate is 1.15
𝑚𝑒𝑎𝑛 𝑆̌
( 𝑡0.90
).

Invasion rate (iα) is nearly constant through time for all α. Extinction rate (eα) is unchanging, species-
charactersitic, and without and additional S-dependent or density-dependent factor included.
Population decline should be most apparent when associated with interaction, especially predation,
yet the small sizes of most populations during the experiment reduced interaction enormously. In
addition, most extinctions during the initial rise of colonization curves are not the result of
interaction, but rather the inability of most species in the Florida Keys pool to colonize these tiny
mangrove islands under any conditions, e.g. lack of proper food or nest site and hostile physical
conditions.
Nagano, T. F. & Burgos, M. G.
Bio 165 – Group 4

Part III: Two-Year Record of Colonization

The time-colonization curves appeared to have assumed the logarithmic forms predicted by basic
equilibrium theory. Moreover, higher levels of species numbers were reached prior to the buildup of
the populations belonging to the constituent species. These numbers then dipped slightly as the
densities of the constituent populations approached the predefaunation levels. The first higher levels
represent “noninteractive species equilibria,” i.e. species equilibria reached before the extinction
rates could be greatly influenced by interspecific interactions such as predation and competition; the
second, lower levels represent “interactive species equilibria,” in which species interactions
contributed significantly to species extinction rates. The equilibrial numbers continue to be an
inverse function of the distance of the islands from the nearest islands/swamps that can serve as
source of immigrants (near island E2 has the most species while distant island E1 has the lowest).
Only E1 has failed to reattain fully the original defaunation level. It is because basic equilibrium
theory predicts that distant islands approach equilibrium more slowly than near islands, and it is
possible that E1 is still in the process of climbing toward equilibrium. Comparison with the data from
the first year of censuses reavealed that a high rate of species turnover occurred during the second
year, resulting in a significant alteration of species composition, even though species number
remained nearly the same.
In time, an “assortative equilibrium” will succeed the “interactive equilibrium” as combinations of
longer-lived species better adapted to the physical conditions of the local environment accumulate.