MALTA'S NATURAL HERITAGE

Full text

Published as:
Schembri, P.J. (1994) Natural heritage. In: Frendo, H. & Friggieri, O. (eds) Malta culture and
identity. pp. 105-124; Valletta, Malta: Ministry of Youth and the Arts; ix + 272pp.
MALTA'S NATURAL HERITAGE
by
Patrick J. Schembri
Department of Biology
University of Malta
Msida, MALTA
INTRODUCTION
Malta
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is justifiably renowned for its rich archaeological, historical and
cultural heritage. However, the islands are generally thought of as having
little to offer in terms of natural history, that is, natural objects and
events, including plant and animal life, fossils, rocks, landscape features
and climate. This is far from the truth. In spite of their small size, the
Maltese Islands are endowed with a variety of habitat types, a diverse fauna
and flora, and an interesting geology. These constitute the natural heritage
of the Maltese Islands, which is just as worthy of study and preservation as
the rest of the nation's patrimony.
Apart from their local importance, some elements of Malta's natural heritage
have a wider regional importance. For example:
 A number of Maltese endemic (i.e. found only in a particular region and
nowhere else) plants and animals are relics from the pre-glacial
Mediterranean flora and fauna and some have no close relatives anywhere else
in the world.
 Numerous endemic species and subspecies of plants and animals have been
described from the Maltese Islands and these are of evolutionary and
biogeographical interest.
 Numerous species of Maltese flora and fauna have a restricted
Mediterranean distribution. Some locally relatively common species are
endangered on a European scale.
 The island of Filfla situated some 4.5km south off the southern coast
of mainland Malta, supports one of the largest known breeding colonies of the
Storm Petrel Hydrobates pelagicus in the Mediterranean.
 The Mediterranean is divided into two major subregions, the East Basin
and the West Basin, each with its own characteristic species. Additionally,
there are differences in species diversity between the northern (European)
shores and the southern (North African) shores of the Mediterranean. Being
situated in the centre of the Mediterranean, the Maltese Islands are at the
meeting point of these four regions, and therefore the marine biota of the
islands is of biogeographical interest.
 The Maltese Islands include the only part of an extensive central
Mediterranean rift system (the Pantelleria Rift) currently to be exposed and
as such provide an insight into the processes associated with development of
this rift.
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In this work, Malta means the Republic of Malta as a whole, while the island of Malta
is referred to as 'mainland Malta'.
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This article briefly reviews the main elements of Malta's natural heritage
and discusses their cultural and scientific importance. There exists an
extensive literature on Maltese natural history, however, there are very few
synthetic works. For reasons of space, it was not possible to include a full
bibliography, therefore, the works listed in the Bibliography were selected
mainly on the basis of their seminality and availability, and on whether they
summarise previous work and include extensive bibliographies. Preference was
given to books and book chapters, however, a fair number of primary research
papers published in learned journals had to be included as, in many cases,
these represent the only available works on the subject.
GEOGRAPHY, GEOLOGY AND PALAEONTOLOGY
Geography
The Maltese Islands are a group of small, low islands aligned in a NW-SE
direction and located in the central Mediterranean at:
latitude: 35°48'28" - 36°05'00" North
longitude: 14°11'04" - 14°34'37" East
They are situated on a shallow shelf, the Malta-Ragusa Rise, part of the
submarine ridge which extends from the Ragusa peninsula of Sicily southwards
to the African coasts of Tunisia and Libya. Geophysically, the Maltese
Islands and the Hyblean Plateau of southeastern Sicily are generally regarded
as forming part of the African continental plate. The islands lie
approximately:
96 km from Sicily (Italy)
290 km from North Africa
1836 km from Gibraltar
1519 km from Alexandria (Egypt)
The sea between the islands and Sicily reaches a maximum depth of not more
than 200m and is mostly less than 90m; that between the islands and North
Africa is much deeper, in places reaching more than 1000m.
The Maltese archipelago consists of three inhabited islands: Malta, Gozo (in
Maltese "Ghawdex"), and Comino (in Maltese "Kemmuna") and a number of small
uninhabited islets: Cominotto (in Maltese "Kemmunett"), Filfola (better known
by its Maltese name "Filfla"), St. Paul's Islands (in Maltese "Il-Gzejjer ta'
San Pawl", also known as Selmunett Islands), Fungus Rock (also known as
General's Rock; in Maltese "Il-Hagra tal-General" or "Il-Gebla tal-General"),
and a few other minor rocks. The land area of the various islands is:
mainland Malta: 245.7 km
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Gozo: 67.1 km
2
Comino: 2.8 km
2
St. Paul's Islands: 10.1 ha
Cominotto: 9.9 ha
Filfla: 2.0 ha
Fungus Rock: 0.7 ha
Geology and palaeontology
Geologically, the islands are composed almost entirely of marine sedimentary
rocks, mainly limestones of Oligo-Miocene age (30-5 million years BP). In
many respects these resemble the mid-Tertiary limestones occurring in the
Ragusa region of Sicily, in the Pelagian Islands, and in the Sirte Basin of
Libya, suggesting that all these localities formed part of the same unit
during their formation. There are also some minor Quaternary deposits of
terrestrial origin. Although the geology of the Malta is overall quite simple
in that it consists only of sedimentary rocks in a basic layer-cake
arrangement, yet in detail it is remarkably complex. Because of the very good
exposures available, the Maltese strata have been the subject of numerous
studies by geologists and are frequently the subject of field-trips organized
by learned societies and educational institutions.
The five main rock types are (in order of decreasing age):
Lower Coralline Limestone which is exposed to a thickness of 140m. This is
the oldest exposed rock type in the Maltese Islands and it started being laid
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down between 30 and 25 million years ago. The coralline algae Lithothamnium
and Archaeolithothamnium (from which the formation gets its name) are locally
abundant, and corals, bivalves and gastropods are characteristic fossils. The
upper part of the formation consists of a c.1m thick bed especially rich in
fossils of the sea urchin Scutella and is thus often referred to as the
Scutella Bed.
Globigerina Limestone is exposed to a thickness ranging from 23m to 207m and
is subdivided into three units (Lower, Middle and Upper Globigerina
Limestones) by two ubiquitous pebble beds. Apart from microfossils (e.g. the
foraminiferan Globigerina from which the formation gets its name), common
fossils of the Globigerina Limestone include bivalves, gastropods and sea
urchins. Remains of turtles, crocodiles, a sirenian, and a seal have also
been found. The pebble beds are rich in fossil bivalves, gastropods, sea
urchins,
corals and shark teeth (e.g. of the giant shark Carcharodon megalodon,
estimated to have attained a length of some 25m).
Blue Clay is exposed to thicknesses of up to 65m. The upper reaches of this
formation are rich in fossils of cephalopods (cuttlefish), bivalves,
gastropods, sea urchins and corals. Vertebrate remains including fish teeth,
cetaceans and sirenians have also been found.
Greensand is exposed to a maximum thickness of 12m, however, in most places
this formation only attains a thickness of some 1m. The name derives from the
abundant dark green grains of the iron mineral glauconite. This horizon is
very rich in fossils, especially foraminifera (e.g. the giant Heterostregina,
sometimes so abundant as to give a 'Heterostregina Limestone'), gastropods,
bivalves and sea urchins.
Upper Coralline Limestone is exposed to a thickness of 162m. This formation
is a complex association of limestones. In some areas, the uppermost parts
show evidence of an intertidal or even supratidal depositional environment
and probably represent the point at which Malta first became dry land, late
in the Miocene, some 10 million years ago.
Localised Quaternary deposits of Pleistocene age (1.9-0.01 million years BP)
occur and comprise 'fossil' soils (palaeosols), fluvial gravels, coastal
conglomerates and breccias, dunes and infillings of caves and fissures.
The more ancient deposits (such as those of the lower beds in the Ghar Dalam
Cave sequence) have yielded the remains of hippopotami (Hippopotamus), dwarf
elephants (Palaeoloxodon), bats, swans (Cygnus) and other birds. Younger
deposits have yielded copious remains of deer (Cervus). Other fossils found
in this and other cave and fissure infills from various parts of the islands
include dormice (Leithia, Eliomys), voles (Pitymys) shrews (Crocidura), bats
(Rhinolophus), an otter (Nesolutra), a bear (Ursus), a fox (Vulpes), a wolf
(Canis), a small horse (Equus) as well as various bird, turtle, toad and
lizard remains.
Many Quaternary deposits, especially the younger red-coloured ones (so
coloured by the oxidation products of iron minerals) are rich in the fossil
and subfossil remains of brackish water, freshwater and terrestrial molluscs.
The Pleistocene sediments and their faunas indicate an overall wetter
climatic regime than that at present. They also indicate that after a brief
period of connection with the Sicilian/Italian mainland, during which there
was an influx of European fauna and flora, the land connection between Malta
and Sicily was severed and the islands underwent a period of isolation
sufficiently long for an endemic island biota to evolve. Much of this biota
became extinct before the end of the Pleistocene, most probably as a result
of changes in the climate. At present there is no very strong evidence for a
land connection with the North African mainland.
Soils
Maltese soils are characterised by their close similarity to the parent rock
material, their relatively young age, the ineffectiveness of the climate in
producing soil horizon development, and the great importance of human
activities in modifying them. Using the Kubiëna classification system,
Maltese soils are of three main types:
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Terra Soils (or Red Mediterranean Soils) which are relic soils formed during
the Pleistocene probably under Mediterranean woodland or scrubland and which
are little affected by the present climate. They are mature and extensively
weathered, have a low calcium carbonate content, and are also low in organic
matter. Terra soils develop on karstland (see below).
Xerorendzinas which are immature soils with a high calcium carbonate content
and low in organic matter. These develop on weathered Globigerina Limestone
and on valley deposits.
Carbonate Raw Soils which are also immature and which have a very high
calcium carbonate content and are very low in organic matter. These develop
on weathered Quaternary sandstones, Greensand, the lower beds of the Upper
Coralline Limestone, Blue Clay and on Globigerina Limestone.
Saline soils and alluvial soils also exist in some areas. In addition there
are soil complexes formed through human agency: either by mixing of powdered
rock with already existing soil at
the time fields were laid out, or by addition of rock debris to soil during
reclamation of disused quarries, or by mixing domestic waste with soil for
use in land reclamation, or by mixing of different soil types transported
from different localities.
Geomorphology
Erosion of the different rock types gives a characteristic topography. Lower
Coralline Limestone forms sheer cliffs which bound the islands to the west;
inland this rock type forms barren grey limestone-platform plateaux on which
karstland develops. Karst is a terrain created by the solution of limestone
rock and is characterised by a series of surface hollows, depressions and
fissures and a subterranean drainage network. The Globigerina Limestone,
which is the most extensive exposed formation, forms a broad rolling
landscape. Blue Clay slumps out from exposed faces to form taluses,
sometimes with slopes of up to 45°, over the underlying rock. Upper Coralline
Limestone forms massive cliffs and limestone-platforms with karstic
topography similar to the Lower Coralline Limestone.
Both main islands are tilted seawards to the northeast. This is interpreted
as being a result of upwelling which started in late Miocene times as a
result of formation of the Pantelleria Rift. There are no mountains, the
highest point is at Ta' Zuta on Dingli Cliffs (SW mainland Malta) which is
253m above sea level; the highest point on Gozo is at Dbiegi (191m). There
are also no lakes, rivers or streams but only minor springs.
The islands are riven by normal faults grouped in two main families: those
trending NE-SW which predominate, and those trending NW-SE. The principal
faults of the NE-SW system are the Great Fault on mainland Malta and the
South Gozo Fault. The Great Fault bisects the island of Malta perpendicular
to its long axis from Fomm ir-Rih on the southwest coast to Madliena on the
northeast coast. In places, the vertical throw of the Great Fault is between
90m and 180m, and produces steep escarpments. The South Gozo Fault runs
parallel to the Great Fault and crosses the island of Gozo from Ras il-Qala
on the east coast to Mgarr ix-Xini on the southeast. Between these two
master faults there is a system of ridges and valleys (see below). The
principal member of the family of NW-SE trending faults is the Maghlaq Fault
along the southern coast of mainland Malta. This fault shows a vertical throw
of some 250m and slickensides (polished and scratched surfaces at the fault
plane produced by friction between the opposing sides of the fault) are very
evident (e.g. at Ix-Xaqqa).
South of the Great Fault, much of the Upper Coralline Limestone, Greensand
and Blue Clay strata have been eroded away, leaving the Globigerina Limestone
exposed. Here, large scale gentle folding is an important structural feature
and this gives southern mainland Malta its characteristic topography of
plains and shallow depressions separated by low hills. South of the Great
Fault it is only in the Rabat-Dingli plateau that all five strata still
remain. Much of the surface of this plateau is typical karstic limestone-
platform.
Block faulting north of the Great Fault gives rise to a sequence of horsts
(ridges) and grabens (valleys) which, proceeding from the Great Fault are:
the Bingemma Basin, Wardija Ridge, Pwales Valley, Bajda Ridge, Mistra Valley,
Mellieha Ridge, Ghadira Valley and Marfa Ridge. The next graben in the
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sequence is inundated by sea water and forms the South Comino Channel separa-
ting mainland Malta from the island of Comino. The highest part of the next
horst is exposed above sea level as the island of Comino and the next graben
is again under water and forms the North Comino Channel.
Topographically, Gozo consists of a series of hills, each topped by an Upper
Coralline Limestone plateau, and separated by low- lying plains where the
rock has been eroded down to the Globigerina stratum. The plateaux are
karstic, the hillsides are covered with clay taluses and the plains between
the hills roll gently.
Characteristic topographic features of particular importance are the rdum,
widien (singular wied), and solution subsidence structures. Rdum are near
vertical faces of rock formed either by erosion or by tectonic movements.
Their bases are invariably surrounded by screes of boulders eroded from the
rdum edges. Because of the shelter they provide and their relative inaccess-
ibility, the rdum sides and boulder screes provide important refuges for many
species of Maltese flora and fauna, including many endemics. Widien are
drainage channels formed either by stream erosion during a previous
(Pleistocene) much wetter climatic regime, or by tectonism, or by a
combination of the two processes. Most widien are now dry valleys, that is,
they only carry water along their watercourses during the wet season; a few
widien drain perennial springs and have some water flowing through them
throughout the year, attaining the character of miniature river valleys. By
virtue of the shelter provided by their sides and their water supply, widien
are one of the richest habitats on the islands; they are also extensively
cultivated.
Changes in sea level have submerged the mouths of some of these widien where
they exit on the coast, giving rise to headlands, creeks and bays. This is
especially evident on the northeastern coasts because of the islands'
seawards tilt in this direction. Especially important are the systems of
drowned valleys which form the creeks of Malta's two main harbours:
Marsamxett Harbour and Grand Harbour, separated by the Valletta headland.
Important examples of inundated river valleys in Gozo include Mgarr ix-Xini
Bay and Xlendi Bay.
A solution subsidence structure results when the surface collapses into a
circular crater-like hollow due to removal by solution of the underlying
limestone. There are two families of such structures in the Maltese Islands:
those formed underwater due to seafloor collapse during the Miocene, and
those formed on land during the Quaternary. The former are only found in
Gozo, and the best examples are Dwejra Bay and Qawra (the 'Inland Sea'). The
terrestrial structures are termed dolines and result from cavern roof
collapse following enlargement of an underground cavern by groundwater. The
best known example of this type of structure is Il-Maqluba on the outskirts
of Qrendi on mainland Malta.
One particular feature of regional importance is the Upper Coralline
Limestone outlier located in the Ghar Lapsi area on the southwestern coast of
mainland Malta. An outlier is an outcrop of rock occurring in a detached
location from the main body of similar rock. In the case of the Ghar Lapsi
outlier, the nearest outcrop of Upper Coralline Limestone occurs some 1km to
the northwest. This outlier is the only part of the extensive Pantelleria
Rift system which is currently exposed above sea level and as such it
provides a unique opportunity for study of the syntectonic depositional
processes associated with rift development; additionally, the younger parts
of the deposit record a Late Miocene emergence of the Maltese Islands better
than that seen in any other Maltese locality.
Climate
The Maltese climate is characterised by moist winters during which the bulk
of the annual rainfall is deposited, air temperatures which never fall below
zero, and a long hot and dry summer. This climatic regime is typical of the
central Mediterranean. The table below gives the mean monthly values of
selected climatic parameters (based on data from the Meteorological Office of
the Department of Civil Aviation).
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Month Rainfall Max. Temp. Min. Temp. Sea Temp. Sunshine
(mm) (°C) (°C) (°C) (h)
Jan 86.4 14.9 10.0 14.5 5.3
Feb 57.7 15.2 10.0 14.5 6.3
Mar 41.8 16.6 10.7 14.5 7.3
Apr 23.2 18.5 12.5 16.1 8.3
May 10.4 22.7 15.6 18.4 10.0
Jun 2.0 27.0 19.2 21.1 11.2
Jul 1.8 29.9 21.9 24.5 12.1
Aug 4.8 30.1 22.5 25.6 11.3
Sep 9.5 27.7 20.9 25.0 8.9
Oct 7.8 23.9 17.7 22.2 7.3
Nov 91.4 20.0 14.4 19.5 6.3
Dec 104.3 16.7 11.4 16.7 5.2
The average annual precipitation is 530mm (mean for period 1951- 1990).
Rainfall is highly variable from year to year; some years are excessively wet
while others are extremely dry (extreme minimum for period 1854-1990,
191.3mm; extreme maximum for period, 1031.2mm). However, no trend in the
annual rainfall appears to exist. The seasonal distribution of rainfall
defines a wet period (October to March with c.85% of the total annual
rainfall) and a dry period (April to September). Even within the wet period,
rain is not evenly distributed. A large amount of rain may fall in a short
period of time during a single storm, and the mean annual rainfall may in
fact represent three or four short torrential downpours. The most intense
outbursts occur during the often violent storms which characterise the
transition from the dry to the wet period.
Air temperatures are moderate (mean annual temperature for period 1951-1990,
18.6°C; mean monthly range, 12.3-26.3°C). The reduced temperature difference
between the warmest and the coldest months of the year is due to the
moderating effect of the sea coupled with the lack of very high ground on the
islands. The islands are small enough for the influence of the sea to be felt
strongly even in inland sites. Temperatures never fall too low for adequate
plant growth. Grass temperatures may fall below zero for a few hours at
night during the period December to April. During the summer months, grass
temperatures may reach values in the upper 40s.
Because of the maritime nature of the islands, relative humidity is
consistently high throughout the year, being mostly in the range 65-80%. The
high relative humidity even during the hot, dry period results in heavy
dewfalls when the temperature falls slightly during the night. These dewfalls
are of extreme importance as they represent the only reliable source of water
for numerous biota during the arid summer months. The Maltese Islands receive
a great deal of sunshine all the year round (mean for period 1951-1990, 8.3h
of bright sunshine per day). The islands are windy, only some 8% of the days
of the year are calm. The predominant wind is the northwesterly which on
average blows on 19% of windy days. The other winds are all nearly equally
represented. Southwesterly winds (known locally as Xlokk) bring damp
oppressive weather and often copious red dust from the Sahara.
The climate of the Maltese Islands has a profound influence on the
vegetation, and consequently, on the islands' fauna, landscape and ecology,
including that of the human population. The hot, arid summer months are very
stressful to plant growth and the natural vegetation is therefore
characterised by evergreen trees and shrubs which resist the adverse summer
heat and drought, and a very large number of herbaceous plants which grow and
flower during the wet period, but which spend the dry period in the form of
seeds or perennating organs (e.g. bulbs, rhizomes etc.) below ground. The
main periods of plant growth are autumn and spring, when temperature and
rainfall are optimal.
FLORA AND FAUNA
The Maltese Islands are popularly regarded as having an impoverished flora
and fauna. This view was probably handed down by casual visitors in colonial
times who, more accustomed to northern latitudes and mainland biotas, drew
wrong conclusions about the totally different environment of the central
Mediterranean. In actual fact, the Maltese Islands harbour a very diverse
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array of plants and animals, especially when considering the relatively small
land area, the limited number of habitat types and the intense human
pressure.
The table below gives estimates of the number of species of selected groups
of plants and animals which occur in the Maltese Islands.
_________________________________________________________________
The number of species of representative groups of plants and animals in the
Maltese Islands. Only terrestrial and freshwater species are considered. In
some cases the numbers given are only estimates as the groups concerned have
not been adequately studied.
_
________________________________________________________________
PLANTS
Algae c.150
Large Fungi c.150
Lichenes (lichens) c.300
Bryophyta (mosses and relatives) c.130
Pteridophyta (ferns and relatives) 11
Gymnospermae (conifers) 2 indigenous species
Angiospermae (flowering plants) c.1000
ANIMALS
Hydrozoa (hydras) 1 species recorded
Turbellaria (flatworms) at least 10 species
Annelida (earthworms and leeches) at least 15 species
Mollusca (snails and slugs) c.67
Arachnida (spiders and relatives) at least 200+ species
Branchiopoda (fairy shrimps, water-fleas
and relatives) at least 10 species
Ostracoda (seed shrimps) at least 7 species
Amphipoda (sandhoppers and beach-hoppers) c.9 recorded
Isopoda (woodlice) c.49
Decapoda (crabs) one freshwater species
Odonata (dragonflies and damselflies) c.10 recorded
Dictyoptera (mantises and cockroaches) c.11
Orthoptera (grasshoppers and relatives) c.48
Coleoptera (beetles) c.600 recorded;
probably 2000+ occur
Heteroptera (true bugs) 113 recorded; more
occur
Lepidoptera (butterflies and moths) c.590
Neuroptera (lacewings) c.12 recorded
Diptera (flies) c.200 recorded;
probably 500+ occur
Hymenoptera (bees, wasps and ants) c.150 recorded;
probably 500+ occur
Diplopoda (millipedes) c.14
Chilopoda (centipedes) c.15
Amphibia (frogs) 1 species
Reptilia (reptiles) 9 species
Aves (birds) c.13 resident (c.57
regular visitors and
c.112 regular migrants)
Mammalia (mammals) c.20 species
________________________________________________________________
As might be expected, the main affinities of the Maltese biota are with
Sicily, the closest landmass of any size. However, the Maltese biota is not
merely an appendage to that of Sicily. To illustrate this let us consider
vascular plants (Tracheophyta) which are much better known than any other
group. The bulk of Maltese vascular plants also occur in Sicily, and indeed
some species are Siculo-Maltese endemics, that is, they are found only in
Sicily and Malta, for example: Sicilian Squill (Scilla sicula), Sicilian Iris
(Iris sicula), Pygmy Groundsel (Senecio pygmaeus), Pignatti's Fern-grass
(Desmazeria pignattii) and Late Spider-orchid (Ophrys oxyrrhyncos).
Nevertheless, several Maltese species are absent from Sicily, for example:
Aleppo Spurge (Euphorbia aleppica) and Olive-leaved Bindweed (Convolvulus
7

oleifolius). The Maltese Islands' position in the centre of the Mediterranean
results in the presence both of western elements such as Mediterranean Willow
(Salix pedicellata), African Tamarisk (Tamarix africana) and Sandarac Gum
Tree (Tetraclinis articulata, the national tree of Malta), and of eastern
elements such as Thorny Burnet (Sarcopoterium spinosum) and Olive-leaved
Bindweed (Convolvulus oleifolius). There is also a fairly strong North
African element represented by such species as Egyptian St.John's Wort
(Triadenia aegyptica), Rock Crosswort (Crucianella rupestris) and perhaps
such Pelago-Maltese endemics (i.e. found only in Malta and the neighbouring
Pelagian Islands) as Cliff Carrot (Daucus rupestris), Maltese Toadflax
(Linaria pseudolaxiflora) and Maltese Waterwort (Elatine gussonei). Similar
patterns of biogeographical affinity are also shown by certain animal groups,
although for these the data are still incomplete.
The Maltese Islands support a number of species of plants and animals which
are found only here and nowhere else in the world. The number of such
endemic species from those groups which have been adequately studied are
given in the table below.
________________________________________________________________
The number of endemic species occurring in the Maltese Islands. Only those
groups which have been adequately studied taxonomically are included and only
freshwater and terrestrial species are considered.
________________________________________________________________
Group Number of endemic species
Tracheophyta (higher plants) 21
Bryophyta (mosses and relatives) 2
Mollusca (snails and slugs) 8
Pseudoscorpiones (false scorpions) 3
Palpigradi (micro-whipscorpions) 1
Isopoda (woodlice) 5
Decapoda (crabs) 1
Thysanura (silverfish) 1
Orthoptera (grasshoppers and relatives) 1
Heteroptera (true bugs) 1
Coleoptera: Staphylinidae (rove beetles) 4
Coleoptera: Elateridae (click beetles) 1
Coleoptera: Tenebrionidae (darkling beetles) 5
Coleoptera: Curculionidae (weevils) 2
Lepidoptera: (butterflies and moths) 17
Hymenoptera: Formicidae (ants) 2
Hymenoptera: Mutillidae (velvet ants) 1
Reptilia (reptiles) 1
Mammalia (mammals) 1
________________________________________________________________
Endemic species are of great cultural and scientific importance. Culturally
they are important because such species are unique to the Maltese Islands and
therefore a valuable part of the national heritage. Scientifically they are
important because of their intrinsic interest with respect to phylogeny,
biogeography and evolution of their group, and for the wider evolutionary
processes they demonstrate. Three cases will suffice to show this: the
endemic vascular plants, the endemic lizard, and the endemic shrew of Gozo.
Some of the endemic plants of the islands are relics from the pre-glacial
Mediterranean flora (these are called palaeoendemics) and have no close
relatives anywhere else in the world. The principal palaeoendemics are
Maltese Cliff-orache (Cremnophyton lanfrancoi), Maltese Rock-centaury
(Palaeocyanus crassifolius, the national plant of Malta), Maltese Salt-tree
(Darniella melitensis), Maltese Fleabane (Chiliadenus bocconei), Maltese Hy-
oseris (Hyoseris frutescens), and Maltese Dwarf Garlic (Allium lojaconei).
The genera Cremnophyton and Palaeocyanus are monotypic, that is, represented
by a single species only, and therefore, these are also endemic to the
Maltese Islands. Palaeocyanus is most closely related to the genus Centaurea
but is more primitive than this and related genera. Cremnophyton is related
to the ancestors of Atriplex. These species are therefore of interest from
the evolutionary point of view since they throw light on the evolution of
certain plant groups. Other endemic plants evolved more recently, following
final separation of the Maltese Islands from the Sicilian and European
mainlands (these are called neoendemics). The neoendemics include Maltese
Sea-lavender (Limonium melitense ), Zerapha's Sea-lavender (Limonium
8

zeraphae), Maltese Pyramidal Orchid (Anacamptis urvilleana), and Maltese Sea-
chamomile (Anthemis urvilleana). These are closely related to mainland
species but differ due to their reproductive isolation. Such species
therefore illustrate evolutionary processes at work.
These processes are also illustrated by the endemic lizard and the shrew of
Gozo. The Maltese Wall Lizard, Podarcis filfolensis, is a species endemic to
the Maltese Islands and the Pelagian Islands of Linosa and Lampione. Four
races have been named from the various islands of the Maltese group
(filfolensis from Filfla; maltensis from mainland Malta, Gozo and Comino;
kieselbachi from St.Paul's Islands; and generalensis from Fungus Rock) and
one race from the Pelagian Islands (laurentiimuelleri). Podarcis filfolensis
is closely related to Podarcis sicula, a southern European species, and to
Podarcis melisellensis, a species of the East Adriatic coast.
Two species of shrew currently occur in the Maltese Islands: the Pygmy Shrew
(Suncus etruscus) which is known to have been introduced into the islands in
historic times and a White-toothed Shrew (genus Crocidura) which is only
found on Gozo and which has been assigned different names by different
workers. Another species of White-toothed Shrew (equated with the extinct
Pleistocene Crocidura esuae of Sicily by some workers) pre-dated human
occupation of the islands. A recent re-evaluation of the living Maltese
Crocidura species has shown that this is actually Crocidura sicula, a
Sicilian species. The present day Sicilian and Maltese species evolved from
the Pleistocene Crocidura esuae and the only real difference between the two
is a reduction in size. Crocidura sicula is a Siculo-Maltese endemism and one
of the few survivors of the Pleistocene fauna of the region. The Gozitan
population has been named as an endemic subspecies, calypso; the other living
subspecies are C.s. sicula of Sicily and C.s. aegatensis of the Egadi
Islands. According to some workers, Crocidura sicula is closely related to
North African species of the genus.
In passing one can mention that many endemic species have been named after
the Maltese naturalists who discovered them. In this way Maltese pioneers of
the study of the natural history of Malta, such as Stefano Zerafa, Gavino
Gulia, Giovanni Gulia, Giuseppe Mamo, Antonio Schembri, Alfredo Caruana
Gatto, Giuseppe Despott, Carmelo De Lucca and Anthony Valletta, are
commemorated.
BIOCOENOSES
Biocoenosis is a general ecological term for any naturally occurring group of
organisms inhabiting a common environment. The terrestrial biocoenoses of the
Maltese Islands may be grouped in two categories: (i) major communities that
are part of the successional sequence towards the climatic climax; and, (ii)
minor communities which are either specialised to occupy particular habitats,
or occupy habitats that are rare in the islands, or are relics from a
previous ecological regime, now surviving in a few refugia. Descriptions of
Maltese biocoenoses are based mainly on vegetation.
 Woodland
It is thought that before man colonised the Maltese Islands, large areas were
covered with Mediterranean Sclerophyll Forest, which is the highest type of
vegetation that can develop in the Mediterranean climate regime. In the
central Mediterranean this forest is characterised by Holm Oak (Quercus ilex)
and Aleppo Pine (Pinus halepensis) with an undergrowth of smaller trees,
shrubs and climbers. The early settlers cut the trees for their wood and to
clear the land for agriculture and buildings. Additionally, these colonisers
introduced sheep and goats to the islands, whose grazing causes some damage
to mature trees but more importantly prevents them from regenerating. In the
Maltese Islands, the native forest is all but extinct and only remnants
remain at four localities, all on the island of Malta. These forest remnants
take the form of small copses of Holm Oak where the total number of trees is
less than thirty. Some of these trees are estimated to be between 500 and
900 years old.
Buskett (mainland Malta) was originally planted by man but is now self-
regenerating and has the character of the natural climax community and may be
described as a semi-natural woodland. Here the wood is dominated by Aleppo
Pine (Pinus halepensis) with various other trees being sub-dominant (e.g.
Olive, Carob, Holm Oak) and there is an extensive undergrowth of shrubs
9

(e.g. Lentisk, Buckthorn and Hawthorn), herbs and climbers. This semi-
natural wood is very important since it represents the only woodland
ecosystem on the islands and consequently harbours a large number of woodland
plants and animals which, because of the lack of suitable habitats in Malta,
are locally very rare. Particularly important woodland species are fungi
which are symbiotic with trees, insects which feed, breed or live in trees
and dead wood, and leaf-litter inhabiting invertebrates.
Many other wooded areas exist in the islands, however, all are man-made (e.g.
public/private gardens, afforestation sites, orchards, etc.) and do not
possess the character of the native climax forest ecosystem nor are they
self-maintaining and self-regenerating, and therefore do not qualify as semi-
natural woodlands.
 Maquis
Maquis is a more or less dense, mostly evergreen shrub community where the
individual shrubs reach a height of between 1m and 3m. In Malta, a semi-
natural maquis develops in relatively inaccessible sites such as the sides of
steep valleys and at the foot of inland cliffs (rdum), while a secondary
maquis develops round trees, mainly olives and carobs, planted by man.
The local maquis is characterised by a number of small shrubs principally
Carob (Ceratonia siliqua), Olive (Olea europaea), Lentisk (Pistacia
lentiscus), Buckthorn (Rhamnus oleoides), Yellow Germander (Teucrium flavum),
Hedge Nettle (Prasium majus) and others.
 Garigue
Garigue is a community of low (less than 1m) scattered, often spiney and
aromatic shrubs with a herbaceous undergrowth. This is the most common
natural vegetation type present in Malta. Some garigue communities are
natural, others result from degradation of forest and maquis. Garigues are
typical of rocky ground, particularly karstland, and are characterised by
such species as Mediterranean Thyme (Coridothymus capitatus), Yellow Kidney-
vetch (Anthyllis hermanniae), Evergreen Germander (Teucrium fructicans),
Mediterranean Heath (Erica multiflora) and the endemic Maltese Spurge
(Euphorbia melitensis), accompanied by numerous geophytes (herbs with
perennating buds below soil level) and therophytes (herbs which survive the
unfavourable season as seeds). Many subtypes of garigue exist.
 Steppic grassland
This is a treeless grassland dominated by grasses, umbellifers, thistles and
geophytes. Steppic grasslands are widespread and result from degradation of
the maquis and garigue, mainly due to grazing (goats are capable of cropping
plants very close to their base thus destroying them and are also able to
chew and eat spiney xerophytic vegetation), but also in response to other
factors. Some steppic communities are, however, climactic or semi-climactic,
for example, those dominated by Esparto Grass (Lygeum spartum) which develop
on clay slopes. The more degraded steppes are characterised by Common Awn-
grass (Stipa capensis), Aegilops (Aegilops geniculata) and a variety of
thistles (e.g. Clustered Carline Thistle Carlina involucrata, Horse Thistle
Notobasis syriaca, Mediterranean Thistle Galactites tomentosa) and geophytes
(e.g. Asphodel Asphodelus aestivuus, Seaside Squill Urginea pancration).
Steppic communities may also develop on abandoned agricultural land.
 Communities of disturbed ground
Given the islands' high human population and its considerable land use, this
biocoenosis has a large coverage. It is dominated by a variety of plant
species, many of which are aliens. Subtypes occur in abandoned fields, along
roadsides and in disturbed seaside habitats.
 Coastal communities
Saline marshlands form an interface between the marine, freshwater and
terrestrial environments. Maltese coastal marshes are characterised by a
muddy substratum on which a pool of brackish water collects in the wet
season. During the dry season this water becomes progressively more brackish
until it finally disappears completely, leaving the marsh dry until the
following wet season. Because of these harsh environmental conditions,
10

saline marshlands support a highly specialised biota which is only found in
this type of habitat. Although several species are common to all local
marshlands, yet each site has its own peculiar habitat characteristics and
suite of species.
Many local sandy beaches were backed by dune systems, but at present only
very few still persist and even these have been much degraded due mainly to
human activities connected with beach development for touristic purposes and
with recreational use. Sand dune ecosystems are thus amongst the rarest and
most threatened of local ecosystems. Local dunes are dominated by the dune
grasses Elymus farctus and Sporobolus arenarius, and, until recently, also by
Southern Marram Grass (Ammophila australis) which has now been totally
extirpated.
On gently sloping rocky shores, halophytic vegetation grows in isolated
patches in the shallow saline soil which accumulates in pockets in the rock.
The species present are typical of this type of habitat and mainly form part
of the Mediterranean vegetational community called the Crithmo-Limonietum. In
the Maltese variants of this biocoenosis, two endemic plants are found only
in this community type and a third also occurs, although it is not exclusive
to low-lying maritime rock. The former are Zerapha's Sea-lavender (Limonium
zeraphae) and Maltese Sea-chamomile (Anthemis urvilleana); the latter is
Maltese Dwarf Garlic (Allium lojaconoi). Other characteristic plants of
scientific importance include: Pignatti's Fern-grass (Desmazeria pignattii)
and Pygmy Groundsel (Senecio pygmaeus), which are Hybleo-Maltese endemics.
Although the fauna of low-lying coastal rock is much less well known, several
species seem to be more or less exclusive to this habitat type.
 Rupestral communities
These grow on cliff faces and high walls. The south, southwest and west
coasts of mainland Malta consist of vertical cliffs rising from the sea to
heights of c.70-130m. In the Dingli Cliffs area, these cliffs give way to a
steeply sloping substratum. This sloping ground is terraced and partly under
cultivation. Further inland there is a second tier of vertical cliffs
(rdum). The south and southwest coasts of Gozo consist of sea-cliffs similar
to those of southern Malta. Because of the shelter they provide and their
relative inaccessibility, both the sea-cliffs, and the second tier of rdum
with the boulder screes which form beneath them, provide important refuges
for many species of Maltese flora and fauna, including many endemics.
The fauna of coastal cliffs includes some of the rarest of Maltese animals;
for example, the endemic Maltese Door-snail (Lampedusa melitensis) occupies a
very precarious habitat of a few tens of square metres only on the southwest
cliffs of mainland Malta, while two other rare endemic snails are found in a
few cliffside localities only. Cliff-side communities are dominated by shrubs
and are especially significant due to the presence of a large number of
endemic plant taxa including the two (Palaeocyanus crassifolius and
Cremnophyton lanfrancoi) belonging to monotypic genera already mentioned.
 Freshwater communities
During the wet season, rainwater collects in natural depressions and hollows
on coralline limestone karstland to form temporary rainwater pools. These
pools are usually very transient and rapidly dry up, especially with onset of
the dry season. These natural freshwater pools house many freshwater species
which are overall rare in the Maltese Islands. A few pools which form in
natural depressions are more or less permanent either because of their
physical size or because they receive water from sources other than
rainwater, usually from springs. These pools are of great local interest
since they represent the only natural standing water bodies in the islands.
Because of the dearth of freshwater in the islands, freshwater plants and
animals are overall rather rare, and this is especially true for those
species which require a more or less year-round supply of water.
The bulk of Maltese plants and animals associating with freshwater are found
in widien watercourses when these are filled with water during the wet
season. One of the most conspicuous species of these habitats is the Painted
Frog (Discoglossus pictus), Malta's only amphibian. Apart from freshwater
species, watercourses support a rich fauna of terrestrial organisms
associated with the luxuriant vegetation that grows along the widien. The
few remaining permanent springs support a distinctive flora and fauna many
11

species of which, since they require a year-round supply of running
freshwater, are limited to this habitat type and are therefore very rare and
have a restricted distribution. One such species is the endemic local race of
the Mediterranean Freshwater Crab (Potamon fluviatile lanfrancoi). Along a
few watercourses there are still remnants of broad-leaved deciduous woodland
with White Poplar (Populus alba), Mediterranean Willow (Salix pedicellata),
and Grey-leaved Elm (Ulmus canescens), sometimes accompanied by Bay Laurel
(Laurus nobilis).
 Caves
In spite of being made up almost exclusively of limestone, the Maltese
Islands have surprisingly few deep caves. It is only recently that the
biology and ecology of Maltese caves has started being investigated. These
studies have shown that Maltese caves are inhabited by organisms which are
adapted to live in such habitats and therefore have a very restricted
distribution. The best known cave-dwellers are bats but there are many other
species, particularly invertebrates. Moreover, a number of these species are
endemic to the Maltese Islands and therefore of great scientific interest.
They are also highly vulnerable, both because of the limited habitat
available and because of their poor dispersive ability. Additionally, many
caves have deposits of Quaternary age, study of which is expected to throw
light on the islands' palaeoenvironment and biogeography. One cave, that at
Ghar Harq Hamiem at St.Georges Bay (mainland Malta) is unique in that it
houses a deep pool of freshwater, the only such body known in the Maltese
Islands.
 Marine biocoenoses
The marine biocoenoses of the Maltese Islands are mostly similar to those
found in other parts of the Mediterranean. Only the most interesting locally
occurring ones of the shore and shallow water off it are discussed below.
Supralittoral biocoenoses are those which occur high up on the shore where
the substratum is only wetted by sea spray and the very highest waves.
Supralittoral communities of rocky substrata are by far the commonest given
that most of the islands' coastline is of this type. Those of soft substrata
are to be found on the few sandy beaches, however, these communities have
been little studied locally. A very distinctive supralittoral biocoenosis is
that of the so called "banquettes", consisting of masses of drying and
decaying plant debris deposited on the shore by wave action. These
specialised communities are composed of an assortment of semi-terrestrial
marine species and semi-aquatic terrestrial species.
Mediolittoral biocoenoses are those which occur in that part of the shore
continuously covered and uncovered by the sea. One such type of biocoenosis
which is considered valuable on a Mediterranean scale and which also occurs
locally, is the vermetid/coralline algal "trottoir" (or 'rim'). This consists
of dense aggregations of vermetid gastropods (Dendropoma sp.) whose uncoiled
shells are cemented to the rocky substratum and to each other, and where the
interstices are filled by the coralline alga Neogoniolithon. A second type of
threatened trottoir, that formed by the coralline alga Lithophyllum
lichenoides may also occur.
Infralittoral biocoenoses are those occurring under the sea, from mean sea
level down to a depth where there is sufficient light for photosynthesis. In
the Maltese Islands this is down to a depth of c.70m. The main community
developing on infralittoral rocky substrata is that dominated by attached
macroalgae. Many subtypes are known, depending on shelter, light penetration,
nature of the substratum, water movement etc. The most widespread are those
dominated by species of the brown seaweed Cystoseira which grow on exposed
rocky shores starting from very shallow water. Another type of Cystoseira
community grows in deep water and is based on the species Cystoseira spinosa,
Cystoseira dubia, and Cystoseira zozteroides. Most of the Mediterranean
species of Cystoseira are endemic to the region.
Sea-grass meadows are perhaps the most important sublittoral biotic
communities in the Mediterranean. Sea-grass meadows are highly productive
ecosystems on which a large number of other ecosystems, and individual
species, depend; for example, many fish and cephalopods use these meadows as
breeding and nursery grounds. Although common and widespread round the
Maltese Islands, sea-grasses are very sensitive to pollution and habitat
12

modification. In many parts of the Mediterranean, these meadows have
regressed and eroded away, leaving in their place much impoverished
thanatocoenoses (i.e. communities based on dead or dying organisms). The same
is likely to happen to Maltese sea-grass meadows in enclosed or semi-enclosed
coastal areas receiving a variety of effluents or subject to certain
activities (for example, dredging, dumping, aquaculture operations etc.).
Several types of sea-grass meadows exist. In deeper water these are formed
mainly by Neptune Grass (Posidonia oceanica), a species endemic to the
Mediterranean. In more sheltered localities and in shallow water, the meadows
are based on Lesser Neptune Grass (Cymodocea nodosa), Red Sea Grass
(Halophila stipulacea) and the alga Caulerpa prolifera.
ACKNOWLEDGEMENTS
I wish to thank Mr Edwin Lanfranco (Department of Biology, University of
Malta) for his help and advice during the preparation of this article,
particularly concerning botanical matters. I thank also Mr John A. Schembri
(Mediterranean Institute, University of Malta), Mr Stephen Schembri
(Mediterranean Institute, University of Malta) and Professor Frank Ventura
(Faculty of Education, University of Malta) for their help with various
aspects of this work, and the Meteorological Office of the Department of
Civil Aviation for supplying meteorological data. I am grateful to the
University of Malta for supporting my research on the faunistics and ecology
of the Maltese Islands.
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