COURSE: ENVIRONMENTAL STUDIES

TEACHER: Dr. DEEPMALA SATPATHY

CLASS: BA PROG SEC-B (TUESDAY- 12.30-01.30AM) (21/04/2020)
Biological Invasion

Introduction Dispersal and movement of species from one place to another is a natural process and
important for the distribution of life in this world. On a longer time scale, we can say that dispersal
is one of driver for evolution of species and responsible for life. Generally, these dispersal and
movement is confined to an area and often limited by multiple barriers such as mountains, rivers,
lakes, oceans and deserts. Species limited to such confined area are known as native species. Every
species take advantage of dispersal and make profit in distribution in range from these processes
to flourishes itself. Man had overcome all these barriers and distributed itself in every part of the
world. The success story of humans in evolutionary time scale is also due to his mental skills and
the capability to domesticate wild species of plants and animals in different region of the world
and utilize them for his benefit. For example, wheat, maize and rice which may be originated and
domesticated in different part of world are now the major cereal crops of the world. There are
hundreds of plant species such as vegetables (potatoes, tomatoes, chilies, brinjal etc.), commercial
crops (groundnut, sugarcane, coffee, tea etc), medicinal plants (cinchona, ephedra, aloe etc) and
animals (sheep, horses, cow, pigs etc) which were always accompanied by man, transported and
introduced from one place to another all around the world for his benefit. Every process has its
pros and cons, while the brighter side is the benefits and economic prospects of the new species
introduced in non-native lands, but darker side is the that some species unintentionally
accompanied with useful species such as weeds, pests, bacteria and viruses. These species created
problems such as epidemics, famines, habitat destruction and homogenization of the biota. Weeds
and pests impacts our agriculture yields while human pathogens can directly linked with humans
and costs millions of valuable lives. Some species escaped from cultivation lands, botanical
gardens and other such human repositories and spread in the nearby forest and other habitat. These
species always accompanied us and move with the human movement to the new places.
Globalization and fast convenient means of transportation with intercontinental connection rapidly
increased the distribution of hundreds and thousands of species intentionally or unintentionally
worldwide. Species introduced to new habitat in a distant land outside their native range separated
through geographical barrier are known as alien species or exotic species or non-native species. In
the medieval period, pathogen Yersinia pestis, which resides in mice and rats causes Plague and
killed about one third of the European population. The potato blight caused by Phytopthora
infestans, a fungus pathogen spread from North America reduced the Irish population to half its
size in 1840. The present times biggest pandemic is AIDS caused by human immunodeficiency
virus spread from central Africa to the rest of the world. At present HIV has infected 1% of the
world population. Besides pathogens, animals like in mammals such as domestic cat (Felis catus),
macaque monkey (Macaca fascicularis), birds such as Indian Myna (Acridotheres tristis), Starlings
(Sturnus vulgaris) and in plants Acacia mearnsii, Lantana camara and Rubus ellepticus are all
included in “100 most worst invasive alien species list” compiled by Global invasive species
programme (GISP). Every year lots countries spend huge amount of money to manage and control
these alien species Biological invasion refers to the disruption of natural communities and
ecosystems by increase in distribution and abundance of alien species outside the native range.
Studies on biological invasion documented the role of alien species in extinction of native species,
disruption of essential ecosystem processes, ill effects on human and animal health and losses to
economy of the region they invade. That’s why it is important to understand alien invasive species
behavior and generate strategies to manage alien invasive species.

Invasion pathways

Introduction of any species to non-native range requires the pathway or means to cross the
geographical barriers such as vast oceans, deserts, rivers and mountain ranges etc. Species
sometime pass these barriers through natural forces like storms, ocean currents or vectors like birds
and animals, but their chances of overcoming these hurdles are meager. Man is one of most
powerful driving force behind all the introduction of species in the non-native range. Throughout
ancient times humans always classified and selected the plant species according to their needs and
exploited them. With the dawn of civilization and advancement in agriculture, trades in-between
the civilization, humans mobilize many alien species to distant lands and continents. In the last
century, with increasing and fast means of traveling, humanaided introduction of species increased
manifold by intentionally and unintentionally means. Many alien species established and
naturalized in the non-native range, but few species spread in vast area and became threat to the
local flora and fauna in that region, these species are known as invasive species. Whereas, the
whole set of processes which results in the introduction of an alien species from one geographic
region to another with vectors including dispersal mechanism and means of introduction is defined
as invasion pathways. The main pathways used by invasive species for their rapidly spread in the
local area are air, rivers, railway tracks, roads, and their dispersal by vectors (man by vehicles and
ships, birds and animals from one region to other). The detailed analysis of human mediated
introduction or pathways includes “causes” for introduction, “vectors” as physical means or
“agents” for transportation, “routes” and “corridors” as geographic paths for transportations.
Horticulturists have always given higher preferences to alien species from native species while
native species are always valued by the ecologists and environmentalists. In case of invasion
pathways, human beings are the prime vector responsible for transportation of invasive species
through vehicles, ships and airplanes others than insects, birds and animals which disperse locally
or occasionally long distances. Routes can be roads, railways, waterways, corridors and air. In
Global invasive species databases (GISD), the invasion pathways were divided into 6 categories:
1) Release (release in nature for use, biological control, erosion control, hunting and fishing in
wild, improvement of landscape/fauna and for conservation and other unintentional release) 2)
Escape (Agriculture, botanical gardens/zoos, farmed animals, forestry, ornamental, horticulture,
research facilities) 3) Transport/contamination (contaminated nursery material, food
contamination, contamination of plants, animals and seeds, trade and transportation of material)
4) Transport (containers, bulk, ship, plane, vehicle, others means of transportation) 5) Corridors
(Interconnected waterways/basins/seas, tunnels and land bridges) 6) Natural dispersal across
borders. The phenomena of different ways invasive species arrive in the non-native range are
known as propagule pressure. The human activities such as intensifying agriculture, clearance of
the forest for building road, railways, urbanizations and over exploitation of native species (timber,
fuel woods, food, fodder, medicines and other commercial activities) are the reasons behind the
spread of alien invasive species. These activities creates disturbance in the natural established
ecosystems hence, creating pathway for the establishment of invasive species into further deep in
the ecosystem. Human induced disturbance was associated with the 97% establishment of invasive
species. Once the species survived in the non native land and starts producing new recruits, it
further spread in the local area by different means of dispersal such as seeds, suckers, roots,
propagules and cuttings. 3.6 Mechanism of Invasion Many theories and framework to explain the
mechanism of the biological invasion which created lot of confusion, hence Blackburn (2011) tried
to unify these all framework (Figure 1). First widely accepted framework for biological invasion
was by Williamson and Fitter (1996) in case of animals and Richardson (2000) in case of plants.
Blackburn took base from Williamson and Fitter (1996) and Richardson (2005) and combine all
other available framework to explain the mechanism of invasion. The invasion framework was the
combination of barriers and stages. There were five vertical bars which signify barriers and four
stages written above the bars in the framework proposed by Richardson. In invasion, the species
has to overcome these barriers to pass into next stage. Blackburn added the additional barrier of
captivity or cultivation after geographical barrier (Figure 1). The arrows signify the species flow
toward next stage and letters in the arrow signify the name of the arrow which is the categorization
scheme for the population. He included the arrow B3 for the many species such as some plants,
fungi, protists, invertebrates, small fishes and other taxa which can skip captivity or cultivation
barrier. The environmental barrier in Richardson’s framework was replaced by survival barrier
which together with reproduction barrier helps in establishment stage. Once the species was
established it remained in C1 and C2 cycle to build enough population (lag period) for dispersal
to long distance. It was very important stage need to overcome by long distance dispersal for
naturalized species to spread and successfully invade. Failure in any stage due to barrier is a failure
for a species to become invader which is shown in the diagram with the horizontal bar and the
arrow. After the alien species had spread, it undergoes through ‘boom and bust’ dynamics which
suggest the increase of the population to its peak and then decline. Terminologies were labeled in
red colour above the diagram whereas management strategies were drawn in yellow arrow at
different stages below the diagram. Conceptualization of invasion is a staged process and not a
linear one. In every stage number of species diminishes explained as invasion failure and filtered
to reach the next stage.

Successful characters of invasive species

It was assumed that invasive species success outside their range is due to some characters which
helped them establish and spread. These traits are as follows:

1. High ecological amplitude i.e. species have larger range of tolerance and can withstands various
environmental condition including long range of temperature, altitude, pH of soil, different terrains
etc e.g. Superior competitor

2. Short juvenile period i.e. starts reproducing from very young age.

3. High fecundity and various mode of reproduction

4. Diverse means of dispersal e.g. disperse seeds through aboitic (wind, air and water) and biotic
means (insects, animals and birds),Long distance dispersal (LDD)
5. Least enemies in the non native range e.g. enemy release hypothesis, and the evolution of
increased competitive ability hypothesis (EICA)

6. Allelopathic effect e.g. Novel Weapon Hypotheses

7. Gaining genetic diversity e.g. through hybridization and founder effects.

Genetic and evolutionary aspect

Researchers believe that the Evolution of increased competitiveness (EICA) helps the invasive
species to shift their resource allocation from the defense against natural enemies in outside their
native range. A small amount of researches had conducted on the evolutionary studies of the
invasive plants and to sort out the answer that certain genotypes within the invading population
are more successful. The ability of exotic plants to establish and rapidly expand in a novel range
is an evolved trait with underlying genetic characteristics in a population. To understand the
genotypes various studies conducted for genetic diversity of invasive species. Genetic variation
occurs due to mutation, selection, gene flow, genetic drift and breeding system in a population
vary over a time. In invasion, ecological adaptation is achieved by exotic species driven by
colonization and subsequent invasion into completely novel range. Local selection occurred from
genetically diverse population which leads to fitness of the species in the new range helps in the
additive genetic variation. Many researchers have found that founder effect associated with the
initial colonization of the species which significantly reduce the genetic variation in invasive
populations. It might reduce their capacity to adapt in novel condition which leads to reduced
genetic diversity due to inbreeding depression limiting propagule production and population
growth whereas other suggest there have been a lag period between initial colonization and
subsequent population of successful invader to rebuild the genetic diversity on which local
population act for selection. Genetic variation is increased in the case of reintroduction of the
invasive plant several times from diverse gene pool of its native range and due to hybridization of
the invasive species with native plant species. However on the bases of the studies on genetic
diversity it was not decided yet that increased genetic diversity leads to invasion because there are
many species which has shown invasive success due to low genetic diversity. It was found that
cross-pollinating plant species have high levels of genetic variation within populations but genetic
differentiation among populations is low, whereas self-pollinating species have less genetic
variation within population and more diversity among populations. Some invasive plants have
attained high genetic diversity within the invaded range such as Phalaris arundinacea, Alliaria
petiolata, Pueraria lobata, and Euphorbia esula where the historical records tell about their multiple
introductions which may be the source of such diversity. Other plant species invaded aggressively
despite having little or no genetic diversity such as Fallopia japonica, Polygonum cuspidatum.
Invasive populations of alligator weed (Alternanthera philoxeroides) and water hyacinth
(Eichhornia crassipes) in China are also reported to have very low levels of genetic diversity.
Reason for these invasions suggested the rapid clonal spread and dispersal via root or stem
fragments in these species for lack of genetic diversity and population genetic structure.
Comparison of the genetic diversity of introduced and native population of invasive plant can
pinpoint which native population is the source of invader and shows the loss of genetic diversity
through founder effect. Diversity data is used to reconstruct invasion histories, reveal how a non
native plant adapts and expands into new territory, can also lead to more effective management
strategies.

Ecological and socio-economic impacts

The impacts of invasive species such as threat to native flora and fauna, disruption of essential
ecosystem processes, ill effects on human and animal health, and losses to economy of the region
they invade is an object of socio-economic interest. Impacts can be measured onto two types: 1)
direct impacts on ecosystem functions and on human welfare, 2) management and control cost of
invasive species plus side effects of introduction of biological control agent. On single species
have varieties of effects on ecology, ecosystem services and socio economic effects e.g. In Cape
region of South Africa, black wattle (Acacia mearnsii) affects the water table, species composition
and also recreational functions due to scarcity of water in rivers and lakes. Ecosystem services
categorized into four main categories are as follows:

1. Supporting services which are necessary for the production of other ecosystem services e.g. Soil
formation, nutrient cycling, Primary production and habitat stability.

2. Provisional services include the production of products obtained from ecosystem services e.g.
Food, fuel, fresh water, genetic resource.
3. Regulating services are benefits supplied by self-maintenance properties of ecosystems e.g.
water regulation, water purification, waste regulation, biological control, pollination, disease
control, natural hazard protection and erosion regulation.

4. Cultural services generate non-material benefits derived from ecosystems e.g. recreational,
aesthetics, education and cultural diversity.

Alien invasive species has been observed to disrupt all these types of ecosystem services which
are directly and indirectly linked with human wellbeing. There are almost 500,000 species of
plants, animals, and microbes have invaded in different countries of the world in which 50,000
have been reported in US alone. However, it is difficult to calculate the economic cost caused by
invasive species. Pimentel (2005) had calculated $120 billion/year USD damage cost from the
invasive alien species in US alone. In Australia the cost estimated to $13.6 billion USD/year and
cost is increasing with each finance year. Most evidence of economic impact of IAS comes from
the developed world. Similarly, there are strong indications that the developing world is
experiencing similar problems due to invasive species. Global invasive species programme (GISP)
was initiated in 1997, internationally to address the growing problems and adverse effect of
invasive species on both natural and managed ecosystems. It had compiled the list of “100 worst
alien invasive species” in its website which had cost trillions of USD damage to the different
countries along with ecological and social impacts. Some of the other important international
initiatives includes Mountain Invasion Research Network (MIREN) and Assessing Large scale
Risks for biodiversity with tested Methods (ALARM) and Delivering Alien Invasive Species
Inventories for Europe (DAISIE) was initiated to address the growing problems and adverse effect
of invasive species on both natural and managed ecosystems in large scale. The ultimate goal of
all these initiative is not only to control and eradicate invasive species but the conservation of
biological diversity, protection of human health and prevention of economic losses. These goals
can be achieved by ecosystem restoration through reintroducing the native species, preservation
of undistributed ecosystem for the establishment of sustainable use by local people. Biological
invasion studies programmes are launched continentally mostly by developed nations for
understanding the behaviors of invasive species and managing them. Since the cost of these
programmes are too much high that developing and underdeveloped world are still not able to
initiate such programmes. It’s still a long way to go, for preserving and restoring the ecosystem by
saving them from biological invasion and restoring the ecosystem services to its normal cycle.