Betonio, Jan Rare R.

ISLAND BIOGEOGRAPHY: HOW THE FLORA AND FAUNA DIVERSIFIED

IN THE PHILIPPINES—A SYNTHESIS PAPER

The answer as to why the island of New Guinea harbours many more species of birds
than on the island of Bali brings us to fact that New Guinea has more than fifty times the area of
Bali and thus, with available space numbers of species ordinarily increases. But why is it that the
island of Hawaii, which is ten times the area of the Louisiades, has fewer native birds? The
theory of island biogeography developed by MacArthur & Wilson has an explanation to such
uneven distribution.

The theory of island biogeography by MacArthur & Wilson has provided a framework in
investigating the ecological and evolutionary dynamics of the processes that control patterns of
species richness. This general theory, with its component equilibrium model, serves as a
structured set of predictions that allows an investigator to formulate biologically important
questions. Systematic approach on the topic of island biogeography allows investigators to judge
their data to similar, comparable data sets, and so may draw conclusions and make predictions.

What is Island Biogeography?

Island biogeography studies the biogeography of the isolated units, the islands, especially
in the context of species diversity and related patterns and ecological processes. Conventionally,
islands are referred to as isolated lands in surrounding waters. Looking in a much broader sense,
‘islands’ also include insular areas or entities such as mountain tops, lakes, oasis, and spring, that
support unique species assemblages relative to surrounding habitats. Oceanic islands, because of
the insular nature, have habitats that are often different from those on the nearest mainland even
when latitudes (climates) and the sizes (areas) are the same. For instance, islands often support
unique species assemblages with more rare and endemic species having small population sizes.
With that being said, islands having the unique features and conservation values are extremely
attractive for intensive efforts in exploration, research, and conservation.

As a major guide in related research ground on islands, MacArthur and Wilson developed
the theory of island biogeography based on observations of many earlier naturalists made during
their explorations around the world. This model has paved the way and continues to inspire many
individuals for further exploration and in some cases has resulted in with much greater effort and
investment in such research.
The Equilibrium Island Biogeography Theory

The ETIB implies that island fauna and flora (biota) eventually reach an equilibrium
point between extinction and immigration. Although species rarely reach equilibrium due to the
extremely dynamic island system, MacArthur and Wilson note that the ETIB permits general
predictions of future island biodiversity patterns.

Apparently, the species–area relationship was discovered much earlier than MacArthur
and Wilson’s island biogeography theory. What is unique development made in the EIBT is the
species-isolation relationship that considers two key ‘island’ features: area and isolation (Figure
1).

Figure 1. A representation of the island biogeography theory that takes both area
(habitat size) and isolation (distance to species sources) into account. The islands
show steeper species–area slopes than the mainland.

MacArthur and Wilson predict that the number of species increases with island area but
decreases with isolation (distance from the mainland or other islands); this prediction laid the
foundation of island biogeography theory. Additionally, the equilibrium island biogeography
theory considers both the immigration (I) and extinction rates (E) at some point are balanced that
is as immigration decreases while extinction rates increases with isolation.

The general island biogeography theory can be considered as two related components: (1)
the effects of area-isolation relationship and (2) the dynamic ‘equilibrium’ between immigration
and extinction. It is therefore assumed, base from the idea of the general patterns in island
biogeography related to area, isolation, and species turnover that at a given time, species richness
on island is static; for a new species to colonize as an immigrant, an existing (resident) species
has to become locally extinct due to competition with other species for niches and resources
(Figure 2).

While MacArthur and Wilson describes immigration and extinction to be symmetrical

across equilibrium for the same island this premise gained several criticism. In fact, most of the
earlier tests of the MW model were based on area and/or isolation only because immigration and
extinction rates need much more time and greater effort to measure. It is likely that the
‘equilibrium’ points (i.e., S1, S2, and S3 in Figure 2), if they ever emerged, would be highly
transitional at best. Yet, it is the core aspect of the MW theory related to ‘equilibrium’ that is still
under debate, especially related to conservation applications.

Island Environment: Factors Affecting the Insular Fauna

The geophysical origins of the island influence the topography of an island. Marine
islands falls into two geophysically distinct categories: continental shelf islands (land-bridge
islands) and oceanic islands. Continental shelf islands are likely to be physically connected to the
mainland during low sea level periods. Due to their connection, these islands have similar
Figure
geological 2. The
structure to theequilibrium island(Williamson
nearby mainland biogeography theory.
1981). ThisS1 , S2,topography,
similar and S3 represent
coupled
equilibrium points of islands with different sizes and distances to the mainland.
with the island’s close proximity to the continent, results in the proliferation of similar flora and
In reality,
fauna. Oceanic I and
islands, E are
on the most
other likely
hand, asymmetrical
are typically in opposite
more isolated, and directions, and their
may have never been
positions and shapes could vary drastically among islands
physically connected to a continental landmass.Island formation therefore, have importantand different
taxonomic groups
implications for island biota because a wide range of elevational gradients brought by this
phenomenon allows for the persistence of diverse habitats.

Moreover, island climate is determined by both external influences, such as ocean

circulation and atmospheric circulation, and internal influences, such as island size, shape, and
topography. If an island is in the path of a moving current or is located where two currents
intersect, this can alter the climate significantly. In addition to circulation influences, the
proximity of an island to a continental landmass also affects the island’s climate. Islands located
close to a mainland, such as land-bridge islands, are likely to be influenced by the continental
climate. Remote oceanic islands, on the contrary, are influenced by the maritime climate. Island
size and elevation, can have a substantial impact on the precipitation. Low islands typically have
relatively dry climates and high islands are wetter and often contain diverse habitats within a
relatively small area. Due to the impact of elevation on island climate, research studies have
indicated that elevation is a critical variable in analyzing species diversity on islands.

BIOGEOGRAPHY OF THE PHILIPPINES

Historical Geology of the Philippines

With the dawn of the field of biogeography, the Philippine archipelago took on an
important role in the development of early biogeographic thought. Over 7000 islands are said to
be enclosed within the boundaries of the Republic of the Philippines, the archipelago containing
islands of widely differing sizes, and geological evidence indicates that several historical groups
exist: some islands are strictly oceanic, some are fragments of once-larger islands, and some had
landbridge connections to the Asian mainland. Wallace used geological explanations (Wallace
1860) to support his discovery of an abrupt faunal transition and considered the Philippines to be
part of the Asian realm. In 1868 Huxley modified the northern portions of Wallace’s Line to
wrap around the Palawan island group, effectively dividing the Philippines between the Asian
region and what would later be known as Wallacea.

The Philippine Islands were formed largely as a result of the collision of a major Pacific
lithospheric plate with the South China Sea. The islands occur as a series of major arcs that are
adjacent to active or inactive subduction zones. Although a small amount of continental crust is
included, it is likely that the islands have formed de novo as an oceanic group.

The islands of the Palawan Arc are combinations of old continental crustal rocks (from
the northern half of Palawan Island to southern Mindoro and northern Panay) and more recent
fragments of oceanic crust. The continental rocks are Palaeozoic and Mesozoic in age and
consist of some clastic sediments and limestone. Sedimentary material of Pliocene and
Quaternary origin on the edge of the Palawan trench suggests the presence of islands by the
Pliocene.

Luzon, the largest and probably

oldest island in the archipelago,
originated as a series of small islands
above water beginning in the late Eocene
or early Oligocene near the Philippine
Trench, and near the Manila Trench by
the early Miocene. By the late Pliocene,
the intervening basin had filled with
enough sediment to project above water,
resulting in a single large

Mindanao, Leyte and Samar are

all associated with the Philippine Trench,
the largest and most active subduction
zone in the area. Southern Mindanao was
also influenced by the Sangihe and
Halmahera arcs; these become more
prominent when they reach the islands of
Celebes and Halmahera. However, much
of Mindanao consists of Pliocene to
recent volcanic deposits, and thus
Mindanao has been a single large island
since the late Pliocene at the earliest.
Leyte consists primarily of Miocene
limestone and volcanics of marine origin.
Samar contains more early Tertiary
volcanic deposits, but also much Figure 3. Pleistocene aggregated islands of the
Miocene-Pliocene limestone. Philippines

The Miocene was marked by migration of the Pacific plate to the northwest over the floor
of the Sulu Sea, forming the Sulu Arc. This subduction zone was active up until the Late
Pleistocene, but is now dormant.

Negros and Panay lie on an active volcanic arc that is bounded on the west by the
actively subducting Negros Trench. Although tectonic activity probably dates from the Miocene,
limestone reefs of Miocene to Pleistocene age are common up to 200 m elevation on many parts
of Negros. Most volcanic materials are of quaternary age. On Panay, sedimentary material is
mostly quaternary in age. Cebu also contains old metamorphosed marine sediments and
volcanics; its time of orgin as an island is uncertain. Camiguin Island, north of central Mindanao,
is composed entirely of quaternary volcanic material from a currently active volcano. Major
eruptions have occurred within historic time.

In summary, although there is evidence that geological activity in the Philippines was
initiated before the Palaeocene, most uplift has occurred since the beginning of the Miocene.
Miocene activity resulted in the formation of at least some subaerial islands, and at least one
large island, Luzon, had formed by the late Pliocene. At the end of the Pliocene much of the
mountain land was reduced to low rolling plains and uplands and, in certain parts at the
beginning of the Pleistocene, these low plains were completely submerged. Much of the present
landform, however, was determined by Plio-Pleistocene tectonic and volcanic activity, and some
areas remain active today.

CHARACTERISTICS OF PHILIPPINE HABITAT

The types of habitat of species influence distribution of species as certain species flourish
only under peculiar conditions. Many species however are not distributed continuously over the
areas which they inhabit, but occur only in suitable stations adapted to their habits and mode of
life. Thus, some will be found only where there are trees, others in the neighborhood of water,
others only on open places.

In the study of distribution of life in the Philippines, the recognition of specific, generic,
and family areas is highly important, as one of the principal objects of this work is to offer an
explanation for at least some of these zoologically and botanically restricted regions, to correlate
them with one another, and to relate them to the neighboring regions.

GENERAL DESCRIPTION OF THE PHILIPPINE FLORA

The Philippine flora is a very rich one,

approximately ten thousand species of flowering plants and
ferns being known from the Archipelago. The flora is in
general essentially Malaysian, its chief alliances being with
that of the Malay Archipelago, but with distinctly
continental (Himalayan) elements in the mountains of
northern Luzon, and a small but interesting series of
Australian types at both low and high altitudes.

The flora is further characterized by a comparatively small percentage of endemic genera

and a high percentage of endemic species, the specific endemism approximating 75 percent.
There is, however, a very striking series of genera and species indicating special alliances with
the islands south and southeast of the Philippines; namely, Celebes, Gilolo, the Moluccas, New
Guinea and, to a certain extent, northeastern Australia and New Caledonia.
In general, the dominant species are identical with widely distributed Indo-Malaysian
species that occur in similar habitats over the vast Indo-Malaysian region. In the primary forests,
however, the dominant species are, for the most part, endemic. The percentage of endemism is
very low in the settled areas at low altitudes, in the open grasslands, and in the secondary forests,
a condition which indicates destruction of the original primary forests and replacement of
endemic by introduced species.

The vegetation of the coastal areas, including the mangrove swamps, is practically
identical with that of similar areas throughout the Malay Archipelago and, for that matter,
throughout the Tropics of the Old World.

The primary forests at low altitudes are characterized by many tree species; between
twenty-five hundred and three thousand are known from the Archipelago. Many of these attain a
large size; especially the dominant Dipterocarpaceae, from which are obtained such commercial
timbers as lauan, apitong, mayapis, yacal, tangile, and others. Gregarious forests (that is, forests
composed chiefly or entirely of a single species) do not exist in the Philippines, except in the
pine regions of Luzon and Mindoro-and there chiefly in the mountains at medium and higher
altitudes-and in the mangrove swamps about mouths of tidal streams. The average primary forest
in the Philippines more closely approximates the mixed deciduous forests of temperate countries,
but is vastly more complex than are such forests. At higher altitudes in the Philippines the
character of the forest changes radically; the species dominant at lower altitudes disappear and
are replaced by representatives of other genera.

The mossy forest, which characterizes the higher mountains of the Philippines, usually
commencing at an altitude of about 800 meters but sometimes not below 1,500 to 1,800 meters,
is composed of many species of trees and shrubs, the trunks and branches of which are covered
with a highly developed moss flora, accompanied by numerous ferns, liverworts, lichens,
epiphytic orchids, and some herbaceous plants.

In the number of known species the family Orchidacea is by far the largest in the
Philippines; nearly nine hundred species of this group are now known. The next largest group is
the Polypodiaceae, a family of ferns; but the entire group of ferns and fern allies includes more
known species in the Philippines than does the family Orchidaceae

SUMMARY OF THE AVIFAUNAL OF THE PHILLIPPINES

  Twenty-five of the seventy avian families
represented in the Philippines are nearly cosmopolitan;
twelve others are represented in four of the zoogeographic
regions of the world but not in the Neotropical or the
Nearctic. The Megapodidse, the Loriidae, and the
Cacatoidoe, each with one Philippine species, indicate an
Australian alliance of the Philippine avifauna. The
Eurylhemidae, with two species; the Trogonidae, with one
species; the Capitonidoe, with two species; the Fringillidse,
with three species; and the Picidae, with twenty species,
indicate a stronger Oriental alliance.

The resident land birds of the Philippines contain a large Austro-Malayan element,
represented by nineteen genera, and a much stronger Indo-Malayan element, represented by
seventy genera. If the Palawan genera that are not represented in the rest of the Philippines be
disregarded, there are still fifty-eight Philippine genera that have little or no Austro-Malayan
alliance. A few genera of the highlands probably came into the Philippines from the north,
possibly through Formosa. Some of the twenty-five endemic genera are unmistakably allied to
Australian genera and others are just as surely Oriental, but most of them are highly specialized
and not very nearly related to extra-Philippine genera.

The Philippine bird fauna includes about seven hundred fifty species in two hundred
ninety-three genera. There is a high percentage of endemism among the species. Few of the
endemic species or genera range over the whole or over a large part of the Archipelago. A study
of the endemic forms leads to the division of the Archipelago into eleven zoogeographic areas,
districts, or subprovinces; these areas are equal neither in size nor in number of restricted bird
forms, and some of them are more clearly cut off than others.

The Palawan group of islands is the most distinct area and exhibits a strong Indo-
Malayan element. The Babuyanes-Batanes area is almost as distinct as the Palawan area, but this
is mainly due to the lack of many of the characteristic Philippine genera and species; this area
contains a small Formosan element.

The Luzon area has the largest number of endemic species and a few of these from the
highlands of northern Luzon are boreal types related to Himalayan genera or identical with them.
Some of these genera are represented in the highlands of Mindanao. The eastern islands; namely,
Luzon, Samar, Leyte, Mindanao, and Basilan, have numerous genera in common that are not
represented in the other islands.
The bird faunae of Luzon, Mindoro, and Palawan are very distinct from each other; the
fauna of Bohol, Cebu, and Negros are also remarkably unlike. The bird fauna of the Romblon
area, as well as that of the Batanes and the Babuyanes, was probably derived from stormdriven
strays or from neighboring areas during brief land connections.

The evidence derived from the distribution of resident Philippine land birds leads to
conclusions, in regard to probable former land connections among the islands and between the
Archipelago and surrounding landmasses that are pleasingly similar in many details to the
deductions from the evidence afforded by geology, hydrography, and phytogeography.

SUMMARY OF THE MAMMALS OF THE PHILIPPINES

From present knowledge of Philippine mammals the following tentative conclusions may
be drawn: The fauna as a whole is dominantly
western Malaysian. The latest invasion came
from Borneo when Palawan and its dependent
islands and the Sulu Archipelago were definitely
land-tied to it. The ancestor of the timarau
(Buzbalus mindorensis Heude) probably reached
Mindoro during the early Pleistocene via a
Palawan peninsula from Borneo. Certain
mammalian elements entered from Celebes upon
a narrow isthmian passage to Mindanao.

Certain elements in the mammalian fauna of northern Luzon may have arrived from New
Guinea upon rafts and succeeded in colonizing. The forms with New Guinean affinities are all
small and hence could make the journey on rafts. The absence of small marsupials, such as
Cuscus, from northern Luzon is puzzling. It is quite probable that such may have landed, but
were exterminated by the more-powerful carnivorous animals already natives of the Philippines.

Formosan mammals may have entered the Philippines from the north during early
Tertiary times by an overland route, but the probability that any Formosan forms drifted by rafts
to the Philippine shores is very slight. An element in the Philippine mammalian fauna showing
distinct affinities with the mammalian fauna of Celebes, New Guinea, and Australia corresponds
to the rather conspicuous floral elements in the Philippine flora. The mammalian fauna of the
Philippines corresponds with the flora in its alliances both with eastern and western Malaysia,
the faunal elements from both regions doubtless having reached the Philippines through channels
similar to those by which the floral elements reached the Islands.

AMPHIBIANS, LIZARDS AND SNAKES OF THE PHILIPPINES

 Most of the Philippine species of amphibians, snakes, and lizards show definite relations
to those of Borneo or the larger Sunda islands. Certain reptilian and amphibian representatives
show relationships with the Moluccas and New Guinea. They are the snake genus Stegonotus;
one lizard of the genus Perochirus, which appears to be confined to Mindanao and the Caroline
Island group; another genus of lizards, Pseudogekko, which has its closest affinity with a New
Guinea genus, Thecadactylus; and another, Otosaurus, whose relation is with Celebes. In the
Amphibia, the three genera Cornufer,
Chaperina, and Phrynixalus show marked relationships
with
New Guinea. The only other genus common to the
Philippines and New Guinea is Rana, of nearly world-
wide
distribution and lacking only in New Zealand.

The order Apoda is composed of a single family,

the Cxeciliidae, represented in the East Indian region by
three genera, which contain four or five species. Of the eleven genera of this family recorded in
Boulenger's Catalogue, the distribution is as follows: Two genera are Malaysian, one of which
occurs in India and Ceylon, and the other in southern Asia and Africa; five genera are American;
two are African; one is American, African, and Asian; one is confined to southern Asia, family
Caeciliide, which restricted to Palawan, Borneo, and the Sulu Archipelago. One of the species
has been found in Basilan. This unique family, with the peculiar biology of one of its species,
Ichthyophis glutinosus, practically requires definite land connections for its dispersal and,
undoubtedly, members of the family spread from Borneo northward into Palawan and the Sulu
Archipelago when the Sulu Archipelago and Palawan were land-tied to Borneo. The amphibian,
reptilian, and saurian faunas of the Islands are still largely unknown and future discoveries will
make changes in the limits of the proposed faunal areas.