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Bicol University College of Science AY 2016-2017 A Partial Fulfillment of Research in Principles of Systematics DIVERSITY ASSESSMENT OF SEAWEEDS AND SEAGRASS IN BARANGAY DIAMANTE, PRIETO DIAZ, SORSOGON Alano, Danielle Daye Almario, Stephanie Arevalo, Angelica Bartolata, Sheena Bola, Immanuel Cañete, Epiphany Ditan, Marc Irvin Nopia, Allaysa Pajarin, Jasmine Rellama, Clarence BS Biology 2A- Group III Submitted to: Prof. Jonathan Jaime G. Guerrero Principles of Systematics DIVERSITY ASSESSMENT OF SEAWEEDS AND SEAGRASS

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Bicol UniversityCollege of Science

AY 2016-2017

A Partial Fulfillment of Research in Principles of Systematics

DIVERSITY ASSESSMENT OF SEAWEEDS AND SEAGRASSIN BARANGAY DIAMANTE, PRIETO DIAZ, SORSOGON

Alano, Danielle DayeAlmario, StephanieArevalo, AngelicaBartolata, Sheena

Bola, ImmanuelCañete, EpiphanyDitan, Marc Irvin

Nopia, AllaysaPajarin, Jasmine

Rellama, Clarence

BS Biology 2A- Group III

Submitted to:

Prof. Jonathan Jaime G. GuerreroPrinciples of Systematics

DIVERSITY ASSESSMENT OF SEAWEEDS AND SEAGRASS

IN BARANGAY DIAMANTE, PRIETO DIAZ, SORSOGON

Chapter IINTRODUCTION

Aquatic life in the Philippines is undeniably diverse. Wide array of

marine organisms ranging from flora to fauna can be found in the

Philippines. Among its areas, Bicol region is one of the well-known provinces

heaving with diverse marine organisms.

The Philippines has the second highest sea grass diversity in the world,

next to Australia. Sea grass meadows provide ecosystem services that rank

among the highest of all ecosystems on earth. It contributes about 19

species or about 55% of the number of species in East Asia. Seven (7)

species, comprising 40% of the total recorded in the Philippines and in

Southeast Asia and 18% of global record, are found in Ulugan Bay in

Palawan.

Seagrasses are a type of submerged aquatic vegetation (SAV) have

evolved from terrestrial plants and have become specialized to live in the

marine environment. Seagrasses possess leaves, roots, conducting tissues,

flowers and seeds, and manufacture their own food via photosynthesis.

Seagrasses support very high biodiversity, and because of their sensitivity to

changes in water quality, they have become recognized as important

indicator species that reflect the overall health of coastal ecosystems. It

provides food, habitat, and nursery areas for a myriad of adult and juvenile

vertebrates and invertebrates. A single acre of seagrass may support as

many as 40,000 fish, and 50 million small invertebrates (Smithsonian Marine

Station, 2002). Sea grasses provide protective shelter for many animals,

including fish, and can also be a direct food source for manatees and

dugongs, turtles, some herbivorous fish and sea urchins. The roots and

rhizomes of sea grasses also stabilize sediments and prevent erosion while

the leaves filter suspended sediments and nutrients from the water column

(Costanza, 1997).

Seaweeds, on the other hand, is the common name for countless

species of marine plants and algae that grow in the ocean as well as in

rivers, lakes, and other water bodies (National Ocean Service, 2014). Some

seaweeds are microscopic, some are enormous but mostly are medium-sized

which come in different colors such as brown, black, red and green. You can

find them randomly wash up on seas and shorelines, or attached to rocks.

Seaweed is packed with a lot of vitamins, minerals, and fiber. Many types of

seaweed do, in fact, possess anti-microbial and anti-inflammatory agents.

Macro algae play important roles in the ecology of coral reefs. They are

the major food source for a wide variety of herbivores and are the basis of

the reef food-web, they are major reef formers, and they create habitat for

invertebrates and vertebrates of ecological and economic importance

(Costanza, 1997). They also play critical roles in reef degradation, when

abundant corals are often replaced by abundant macro algae. This may

result from over-fishing of herbivorous fish or from pollution by excess

nutrients and sediments. Increased macro algae on a coral reef are often

undesirable, indicating reef degradation, although this depends on the type

of algae (Duarte, 1999).

Seaweeds are a good source of colloidal materials which are used as

gelling agents, emulsifiers, stabilizers, in pharmaceutical, cosmetic and food

products. Seaweeds also constitute an important food item, fertilizer and

animal feed.(Legasto, R.)

Statement of the Problem

The abundance of seaweeds and seagrasses inevitably brings about

negligence and passivity of people in taking care of these marine organisms.

There is complacency in recognizing its importance which leads to inactive

utilization or improper treatment. Awareness of these organisms must be

raised in order to locally promote better conservation and consumption. For

optimal preservation, people must have a basic knowledge about these

organisms. With such concern, a cohesive research regarding the diversity

assessment of seaweeds and seagrasses in Barangay Diamante, Prieto Diaz,

Sorsogon will be conducted.

Objectives of the Study

Generally, the study aims to identify and assess the seaweeds and

seagrasses present in Barangay Diamante, Prieto Diaz, Sorsogon.

Specifically, the study will pursue the following objectives:

1. Identify the species of seaweeds and seagrasses present in the

selected study site

2. Assess the community structures of seaweeds and seagrasses in

the area in terms of:

a. Frequencyb. Densityc. Biomass

3. Determine the diversity indices of seaweeds and seagrasses

present in the area

a. Dominanceb. Evenness

4. Construct a dichotomous key in order to identify the species of

seaweeds and seagrasses based upon their characteristics or

morphological features

Significance of the Study

This study aims to help further apprehend the importance of sea

grasses and seaweeds in the marine ecosystem in Brgy. Diamante, Prieto

Diaz, Sorsogon. By means of studying and focusing on the: (1) Identification

of species of seaweeds and sea grasses present in the selected study site,

(2) Assessment of the community structures of seaweeds and seagrasses in

the area in terms of: frequency, density, and biomass, (3) Determination of

the diversity indices of seaweeds and sea grasses present in the area and (4)

Construction of dichotomous key in order to identify the species of seaweeds

and sea grasses based upon their characteristics or morphological features.

This research will also contemporize to the previous studies that will

raise the awareness of residents and local groups of Department of

Environment and Natural Resources (DENR) with the cohesive updates and

recent assessment regarding these organisms.

Chapter II

REVIEW OF RELATED STUDIES AND LITERATURE

This chapter comprises the related studies and literature from cohesive

researches related to the attainment of this study’s objectives.

Queding (2011) in the sea grass and seaweeds of Cagmanaba, Oas,

Albay using the transect line method in the 3 stations, the seaweeds and sea

grass collected were assessed. In the results obtained,3 species of sea grass

belong to two (2) families, the Enhalusacoroides and Halophilic minor in

family Hydrocharitaceae and Halophilic minor in family

Potamogetonaceae .About the seaweeds, six (6) seaweeds were found, four

(4) in the Division Phaeophyta, one (1) in Division Chlorophyta and one (1) in

Division Rhodophyta. The most dominant in the sea grass was

Enhalusacoroides and Padina minor in the seaweeds community.

In the assessment of Sta. Cruz (2011), in the sea grass and seaweeds

of Buenavista, Legazpi City using the transect line method in 3 sampling

sites, he found out that there was an abundance of seaweeds and sea

grasses in the area. Two thousand five hundred and three (2,503) seaweeds

and one hundred forty six (146) sea grasses were collected in the 3 sites.

There were 4 species of sea grasses that belong to two families of the

Enhalusacoroides in family Hydrocharitaceae and Cynodoceaserrulata,

Halodulepinifolia and Haloduleuninervis in family Cymodoceaceae. The

Cymodoceaserrulata was the most abundant while the Halodulepinifolia was

the least. Sta. Cruz also asserted that the use of fish rod and fish net was

more efficient compared to scissors net and trawls and active fishing gears

which are being used in the Buenavista.

In the study conducted by Lobigan (2011) in the seaweeds and sea

grass of BarangayPantao, Libon, Albay using the transect line method in 3

sampling stations, he assessed the seaweeds and sea grasses that were

found and in this assessment the results that was obtained is that there were

2 species of sea grass found, the Halonduleunivervis in family

Potamogetonacea and Halophila minor in family Hydrocharitacea. In the said

species, the Halophila minor was more dominant in the two sampling

stations. Lobigan also said that having a proper knowledge of the public

about the state of sea grass can help in the conservation of the said species.

Taxonomic and ecological studies on Philippine seaweeds have been

conducted. In Northwestern Luzon, the earliest studies were conducted by

Domantay (1961) and Galutira and Velasquez (1963). Domantay reported

the algal vegetation of Hundred Islands, Pangasinan. Galutira and Velasquez

identified some species of edible seaweeds in Ilocos Norte. The species

included Porphyra, Gracilaria, Hypnea, Acanthophora, Caulerpa, and Codium

among others. In 1979, Moreland made a study on the edible seaweeds of

Northern Luzon. His study focused on the market prices, local taste

preference, seaweed recipes and other local uses. No mention was made on

the spatial and seasonal distribution of these resources.

The natural tendency of Filipinos to locate themselves near or along

the coastal areas and the archipelagic nature of the country have led to the

very close dependence of the coastal inhabitants on the sea as a source of

nutriment and livelihood (Trono, 1988). Seaweeds and seaweed products are

among the resources on which the lives of coastal inhabitants are very much

dependent.

SEAWEEDS

Seaweeds are large, attached forms of marine algae that

show high diversity in rocky shores, where they are often subjected to tides

and waves. They comprise more than 90% of all marine plants (Ismail,1995).

Seaweed can grow as individuals, but they move more frequently

live together in communities with other seaweed and animal species.

Seaweed communities affect and are affected by the environment and are

some of the most productive marine plant communities in the world (Dawes

1998).

Seaweeds are large algae (macro algae) that grow in salt water or

marine environment and lack true stems, roots and leaves (Krishnamurthy,

1996). Seaweeds commonly grow on coral reefs or in rocky landscape or can

grow at great depths if sunlight can penetrate through the water (Mateljan,

2006). Most of the seaweeds can be seen thriving in underwater beds

floating along the sea surface attached to rocks (Collen, 2001). Seaweeds

play an important and vital role in the marine ecosystem, providing food and

shelter for host of creatures, such as sea urchins, lobsters and young fishes.

`Macroalgae, or seaweeds; are multicellular, macroscopic, marine

algae defined as nonvascular plants. They are cultivated near-shore at

shallow depth, where it can attach to bottom substrates ( natural or artificial;

seaweeds cannot grow in sandy bottoms), or at greater depth, where

underwater ropes, strung out in long lines anchor seaweeds to the bottom ,

typically in areas protected from direct stoorm surge (Rothe, Hays,

Benemann,2012).

Seaweeds are macrobenthic algae. It is composed of three Phylum;

Chlorophyta which means green plant. Examples of this are, Ulva, Caulerpa,

Halimeda and Codium. Phylum Phaeophyta, thus phylum refers to brown

algae, unlike the green and red algae that can live both in saltwater and

freshwater, brown algae all of which are marine. This includes the largest

algae- the kelp. Giant kelps grow to a length of more than 90 meters. There

are no kelps in the Philippines. Examples of brown algae are, kelps,

sargassum and laminaria. Porphyra a red alga, belongs to Phylum

Rhodophyta which means red plant most of which are marine; only 2% are

freshwater species. Example of this is the Corallina which has tiny hard

segments. Like the green alga Halimeda, it also contains calcium carbonate.

Segments of three two kinds of algae get piled up among corals, that

contribute to the formation of coral reefs (Joaquin C.C , Rabago L., Lagunzad

C. 1996).

Seaweeds Diversity and Distribution

Some 820 species of marine macrobenthic algae, including many

species of Cyanophyta, are reported from the Philippines. These consist of

472 species of Rhodophyta belonging to 37 families and 11 orders. 134

species of Phaeophyta belonging to 10 families and 7 orders and 214 species

of Chlorophyta belonging to 11 families and 7 orders. The Rhodophyta

comprise 57.6% the Phaeophyta 16.3% and Chlorophyta 26.1% of the flora.

Many of these species are of economic importance as food, sources of

industrial products such as polysaccharides, bioactive and nutritional natural

products, and growth promoting substance (Trono, 1999).

Seaweeds are large, attached forms of marine algae that show high

diversity in rocky shores, where they are often subjected to tides and waves.

They comprise more than 90% of all marine plants (Ismail, 1995) and are the

most important primary producers in the marine ecosystem (Trono, 1998).

Studies on the diversity of seaweeds had been conducted around South-east

Asia in countries such as the Philippines (Trono, 1999), Singapore (Lee et al.,

2009) and Thailand (Prathep et al., 2011; Ruangchuay et al., 2011). In

Malaysia, studies on the composition and diversity of seaweeds had been

conducted mainly in Peninsular Malaysia by Ismail (1995), Phang et al.

(2005), Phang (2006), Phang et al. (2008) and Gan et al. (2011). There is

hardly any study on seaweed diversity in East Malaysia.

Seaweeds Classification

The classification of algae is complex and somewhat controversial, especially concerning the blue-green algae (Cyanobacteria), which are sometimes known as blue-green bacteria or Cyanophyta and sometimes included in the Chlorophyta. The following provides a taxonomical outline of algae.

Archaeplastida

•Chlorophyta (green algae)

• Rhodophyta (red algae)

• Glaucophyta Rhizaria, Excavata

• Chlorarachniophytes

Algae and aquatic macrophytes as feed in small-scale aquaculture

• Euglenids Chromista, Alveolata

• Heterokonts

• Bacillariophyceae (diatoms)

• Axodine

• Bolidomonas

• Eustigmatophyceae

• Phaeophyceae (brown algae)

• Chrysophyceae (golden algae)

• Raphidophyceae

• Synurophyceae

• Xanthophyceae (yellow-green algae)

• Cryptophyta

• Dinoflagellates

• Haptophyta

Importance of seaweeds

Seaweeds, or macroalgae are an ecologically and economically

important component of marine ecosystem worldwide. They are primary

producers, shelter, nursery grounds and food sources for marine organisms.

In addition, they are used around the world as foods and fertilizers, and for

the extraction of valuable commercial products, such as the industrial gums

and chemicals ( agars, carageenans, and alginates) (Prathep, 2005).

Seaweeds were increasingly used in the industrial, scientific, chemical

plant fertilizer and as livestock feed supplement (Druehl, 2000). Several

seaweed species have economic importance as food for humans, as

industrial material and as ingredients in traditional medicine (Hong et al.,

2007)

Seaweeds are extensively used as food by coastal peoples all over the

world. Red algae like porphyra and nori are used in soups and salads. Irish

moss is another red alga used in producing various food additives along with

Kappaphycus. Alginate, agar and carrageenan gelatinous substances

collectively known as hydrocolloids or phycocolloids. Hydrocolloids have

attained commercial significance especially in food production. Most of the

carrageenan is extracted from Kappaphycusalvarezii and

Euchemadenticultum. The original source of carrageenan was

Chondruscrispus. Carrageenan was a family of linear sulphated

polysaccharide extracted from red seaweed. The name is derived from a

type of seaweed that is abundant along the Irish coast line (Buck et al.,

2006).

SEAGRASSES

Seagrass are angiosperms ( flowering plants) more closely related to

terrestrial lilies and gingers than to true grasses. They grow in sediment on

the sea floor with erect, elongate leaves and a buried root-like structure

(rhizome).

Seagrass are specialized marine flowering plants that have adapted to

the near shore environment of most of the world’s continents (Short et al.,

2001), except Antarctica (den Hartog, 1970).

Seagrasses are underwater flowering plants that evolved from land

plants around 100 million years ago. Seagrasses grow on sandy to muddy

substrates, from the intertidal zone to a depth of approximately 40 metres.

They are anchored to the substrate by a network of fibrous underground

stems or rhizomes. Their distribution is dependent upon temperature and

exposure to wave action but, most importantly, upon light. The depth

distribution of seagrasses is therefore influenced by water clarity, as this

controls light penetration. In South Australia, seagrass meadows cover

approximately 5000 km 2 of sea floor (Shepherd & Robertson 1989).

Seagrass plants generally shed their leaves annually in early autumn; the

blades are initially buoyant, then sink and accumulate in drifts on the

seabed. Nutrient levels in the surrounding waters have been shown to play a

role in leaf shedding as well. During periods of strong wind and wave activity,

particularly in the winter months, surge and swell resuspends and mobilizes

the detached leaf blades. Persistent onshore winds blow this drift-cast

material into the surf zone and at high tide it is washed onto the beach in

clumps. Greater than 95% of the beach-cast seagrass in South Australia

consists of the blades of Posidonia spp. (Management Plan 2000).

The Historical account of seagrass research in the Philippines includes

an advancement in the effort of more than four decades of scientific

research in the Philippines. The studies started with the taxonomy,

distribution and structural accounts of selected species, and continuing at

present, with ecology, physiology, metabolism, and responses to

environmental and climate change impacts. The shifting needs of the times

aggravated, principally by an alarming reduction and loss in resources

resulting from a decline in coastal water quality and degradation of the

environment, dictating a corresponding shift in seagrass research focus from

basic to its application, from purely scientific initiatives to those that now

require support and collaboration from social and behavioral sciences.

(Marine Science Institute, Fortes 2003).

Seagrass are prominent component of Philippine coastal ecosystems

(Fortes 1995, cited by Genito, Nabuab, Acabado, Albasin and Beldia) they

are biota and habitat in one, as they naturally and simultaneously function

both as primary producers and structural species. Consequently, they form

ecosystems of great physical, biological, and economic importance (Baron, et

all., 1993) as they sustain a diversity of fauna, act as nutrient sinks and

sedimentation buffers, and generally support important human exploitation (

Hirst and Atrill,2008; van Houte-Howes et all.,2004; Paula et all.,2001).

Moreover precisely because they are ecological constant of shallow coastal

areas, they are rendered vulnerable to habitat fragmentation and sediment

loading mostly brought about anthropogenic activities (Hirst and

Atrill,2008;Tanner,2005; Fortes and Santos,2004).

Seagrass Diversity and Distribution

Eighteen seagrass species were found from 529 sites in the Philippines.

In relation to seagrass as a resource in need of protection, its status as such

is yet largely unknown, becoming a focus of scientific inquiry only in the last

30 years and, and as an object of conservation, only in the last 15 years. The

coastal nature of Philippine demography, in addition to numerous

development facilities, has caused eutrophication of marine waters, which,

along with habitat loss, is a major long-term threat to seagrass ecosystems.

Some advancements in seagrass research were made locally that are useful

steps to reverse seagrass habitat loss. These steps include (1) focusing

research on management issues; (2) developing an integrated framework for

action; (3) undertaking an economic valuation of the resource; (4) using

available scientific knowledge as a means to forge public-private

partnerships; (5) ensuring a functional coordination among concerned

agencies; and (6) ensuring high quality scientific publications. (Fortes M,

2003)

There are 58-60 describe species of seagrass worldwide, within 12

genera, 4 families and orders. The Indo-Pacific has the largest number or

seagrass species worldwide, with vast meadows of mixed species stands.

There are 23 species(Short et al.,2001) of seagrasses found throughout the

tropical Indo-Pacific (Region IX, in Short and Coles 2001). These includes the

genera of Cymodocea, Enhalus, Halodule, Halophila, Syringodium, Thalassia,

Zostera and Thalassodendron. Seagrasses provide a sheltered, nutrient-rich

habitat for a diverse range of flora and fauna. The Philippines is believed to

be the area where seagrasses originally evolved, and has a high

concentration of seagrass species. In the Western Pacific there are 16

species recorded from the Philippines, 13 from Papua New Guinea,

( Fortes,1998), and 15 from Northern Australia ( Lee Long et al., 2000).

Tropical seagrasses occupy a variety of coastal habitats. Tropical

seagrass meadows typically occur in most shallow, sheltered soft-bottomed

marine coastlines and estuaries. These meadows may be monospecific or

may consist of multi species communities, sometimes up to 12 species

present within one location. The depth range of seagrass is usually control at

its deepest edge by the availability of light for photosynthesis. Most tropical

species are found in water less than 10 m deep. Of the 13 species identified

in Northern Queensland by Lee Long et al.,(1993) most occurred in water

depths less than 6m below mean sea level(MSL) and only four occurred in

water more than 20m below MSL. Coles et al.,(1987) noted three general

depth zones of seagrass species composition for tropical waters; a shallow

zones less than 6m deep with high species diversity include all species found

in a region: a zone between 6 and 11m most commonly found were the

Halodule and Halophila species: and a zone of deeper than 11m where only

species of the genus Halophila were found.

Species of the genus Halophila are common throughout the tropics

and can be found in a range of habitat types form shallow estuarine

environments to very deep clear water. Example, H.decipiens grows to 58m

in the Great Barrier Reef ( Lee Long et al.,1996) and H.spinulosa, H.ovalis,

H.tricostata, and H.capricomiwere common below 35m (Coles et al.,2000).

Halophilaovalis is probably the most widely distributed tropical seagrass

species, occupying a wide depth range in the Indian and Pacific oceans.

Thalassiahemprichiiis often associated with coral reefs and is common on

reef platforms where it may form dense meadows, it can also found

colonizing muddy substrates, particularly where water pools at low

tide(McKenzie L.J. and Campbell S.J., 2002).

Importance of seagrasses

Seagrass on the other hand, are marine angiosperms that are adapted

to live permanently in drenched seawater. Worldwide, there are about 58

species.(Bandeira, 1995; Richmond, 1997).

Seagrass provide food and habitat to numerous marine species,

stabilize the ocean button, maintain water quality , and help support local

economies (Jackson et al.,2001, McManus and, Polsenberg 2004, Orth et

all.,2006).

Seagrasses are marine angiosperms that occupy shallow nearshore environments. Seagrasses provide important ecological, social and commercial functions. They act as ‘ecosystem engineers’, stabilizing sediments and attenuating water flow, and as bio-indicators or ‘coastal canaries’. Seagrasses support high biodiversity and provide habitat for species of recreational and commercial value. Seagrasses also perform important ecosystem functions, including trophic subsidies and carbon storage (F. Y. Warry, J. S. Hindell 2009).

Chapter III

METHODOLOGY

The research will be conducted for three (3) days, from August 19 to

21, 2016. The researchers will be divided in groups to maximize the time of

assessment. Each sampling will be done for few hours in every sampling area

depending on the increasing tides.

Study Site

The study will be conducted at Prieto Diaz which is 387 km (241mi)

southeast of Manila, 50 km (31m) east-southeast of Legazpi City the regional

center and 20km (12.4mi) north-northeast of Sorsogon City the provincial

capital. North of Prieto Diaz across the Albay Gulf is Rapu-Rapu, Albay, on

the west is Bacon District of Sorsogon City and on the south is Gubat,

Sorsogon while Pacific Ocean is on the east.

Prieto Diaz has a tropical climate. There is significant rainfall

throughout the year in Prieto Diaz. Even the driest month still has a lot of

rainfall. According to Köppen and Geiger climate is classified as Af. The

average annual temperature in Prieto Diaz is 27.1 °C.

Vicinity map of Prieto Diaz, Sorsogon

Sampling Technique

Line quadrat method, which is a standard method in the analysis of plant

communities, will be used. Three (3) 50 meter transect lines will be laid at

the chosen areas, perpendicular to the shoreline. The 0.5 X 0.5 m quadrat

with 25 sub quadrats will be randomly placed on each side of the transect

line. Then, the seagrasses and seaweeds will be identified, counted and

estimated for their frequency and cover.

Identification of Seagrass and Seaweeds

After sampling, the harvested samples of seagrass and seaweed will be

rinsed, and cleared from adhering debris. Then, the samples will be placed in

clean plastic bags for biomass evaluation by weighing the labeled samples.

Identification of species will be done using some taxonomic keys and

references and with the guidance of an expertbased on the observed

morphological appearance.

The Line Transect Method

Preserving of the Specimens

Collected specimens will be carefully washed to remove the sand from

the ocean and the epiphytes attached to it. Each species will be placed in

different zip locks. They will be soaked in ethanol to be preserved. After

soaking the for a few days, they were pressed with newspapers

Community structure analysis

Adapting previous and similar studies, the seaweed and seagrass

species that will be collected in line transect method will be described using

the following formula. This is in line with the attainment of the objectives.

Species density= Number of individual speciesTotalarea sampled

Relativedensity=Density of individual speciesTotal density of all species

x100%

Ecological Indices

Shannon’s index of general diversity (h’) will be used to determine the

diversity of species in the study site.

H’= -Pi (In Pi)

Where:

Pi = importance probability of each species (ni/N)

Ni = number of individuals in each species

N = total number of individuals in all species

In = natural logarithm

Index of dominance(C) will also be used to compare the dominance

of the same species in an area.

∁=( ¿N

)2

Where:

N = total number of individuals of all species

ni = total number of individuals of species A

Evenness is important to determine the distribution of species in a

certain area.

ϵ = H'

logS

Where:

H’ = value of Shannon’s General Diversity Index

S = total number of species