studies on eco-biology of some freshwater crabs from jammu...

Studies on Eco-biology of some freshwater crabs from

Jammu

Thesis

Submitted to the University of Jammu

for the award of Degree

of

Doctor of Philosophy

in

Zoology

By:

Meenakshi Bandral

Under the Supervision of

Prof. Kadambari Gupta

Prof. Seema Langer

Post Graduate Department of Zoology,

University of Jammu, Jammu- 180006

2015

P.G. DEPARTMENT OF ZOOLOGY

UNIVERSITY OF JAMMU, JAMMU - 180006

No. JU/ ZOOLOGY/…. Dated: …………….

CERTIFICATE

1. Certified that the thesis entitled, “Studies on Eco-biology of some freshwater

crabs from Jammu” embodies the work done by the candidate Mrs. Meenakshi

Bandral.

2. The candidate has worked under our supervision for the period required under

rules.

3. The candidate has put in the required attendance in the department during the

period required under statutes.

4. The thesis being submitted for the degree of Ph.D. in Zoology is worthy of

consideration for the award of Ph.D. Degree by the University of Jammu, Jammu.

5. The conduct and character of the candidate remained excellent during the period

of research.

Prof. Kadambari Gupta (Supervisor)

Prof. Roopma Gandotra

(Head of the Department)

Prof. Seema Langer

(Supervisor)

TO MY GUIDES

Words cannot describe my heartfelt gratitude and appreciation for my

research supervisors, Prof. Kadambari Gupta and Prof. Seema Langer,

Department of Zoology, University of Jammu for their encouragement and

moral support. I sincerely thank them for their advice, wise words and help

at every point of time during the period of research. Their positive approach

and enthusiasm constantly instilled in me, the crave to learn more and more.

Their effective directions, scholarly guidance, inspiring and valuable

suggestions helped me to compile this thesis. They always made themself

available to me for the checking of the manuscript and rendering

constructive additions in it. They are true mentors and persons of great

understanding, perseverance, immense knowledge and patience. I feel myself

honoured and fortunate to have being worked with such wonderful guides

and above all, great human beings. The respect they owe is far more than

that one can express in words.

Acknowledgements

ACKNOWLEDGEMENTS

I thank the almighty for his benign presence with me all through the journey of the

preparation of this thesis.

I would like to express my warm and sincere thanks to Professor Roopma Gandotra,

Head of the Department, University of Jammu, Jammu for the blessings and support

besides providing me laboratory and library facilities to carry out my research work

smoothly. I wish to extend my deep sense of gratitude to Prof. K.K. Sharma (Ex-HoD)

for his encouragement and unflinching support during his headship period.

I am highly indebted to all the learned members of the Teaching faculty of the

department, Prof. D.N. Sahi, Prof. N. K. Tirpathi, Prof. J. S. Tara, Dr. Sanjay Bhatia,

Dr. Arti Sharma, Dr. Parvinder Kumar, Dr. Sarbjeet for their timely support and help.

I owe my thanks to the non-teaching staff of the Deptt of Zoology, for their kind help,

support and cooperation.

My sincere thanks to Madam Kiran Bakshi (Ex-Principal G.C.W, Parade) for her kind

cooperation while seeking permission from the higher authorities to undertake this

research project. The perusal of help and support rendered by the non teaching staff of

G.C.W. Parade under the able guidance of Madam Hemla Aggarwal (Principal G.C.W.

Parade) is also acknowleged.

I am highly thankful to the Department of Higher Education, Government of J&K for

allowing me to undertake this research work.

I wish to place on record my special and heartfelt thanks to my Lab. Mates, Mr.

Rakesh Gupta, Ms. Priya Manhas, Ms. Bipu Khajuria, Ms. Arti Devi, Ms. Vipulab

Sharma, Ms. Neha Anthal, Mr. Nipun Sharma and Ms. Chhavi Sharma, for their

cooperation , help and support.

I owe a deep sense of gratitude to my Father Sh. K.S. Bandral, Mother Smt. Raj

Bandral and Father in Law Sh. C.S. Chib Mother in law Smt. Padma Chib whose

blessings served as my success keys to complete my research project.

Special thanks to Mr. K.S. Chib (KAS) my soul mate, for his constant support, patience

and love. He has been the driving force and inspiration behind this work. This piece of

work would have not been complete without his encouragement, unconditional support

and help.

The affection and love shown by my daughters- Sunidhi and Manya Chib helped me to

sway through the difficult moments as I took their precious moments when they needed

me the most.

Brotherly help of Lt. Col. Vikram Khajuria is greatly acknowledged in facilitating in

approaching ZSI (Zoological Survey of India) Kolkatta and help rendered by M.K. Dev

Roy and Shantanu Mitra for identification of the specimens is also highly

acknowledged. Timely assistance rendered by Mr. Ankur Kumar Sharma is highly

appreciated in fulfilling my endeavourer with zeal and devotion.

The financial assistance provided by UGC ,New Delhi, FIP (Fellowship Improvement

Programme) is also greatly acknowledged.

Last but not least, all may not have been mentioned but none is forgotten.

Meenakshi Bandral

CONTENTS

Chapters___________________________ ________________________________________

Acknowledgements

1. Introduction 1-6

2. Review of Literature 7-42

3. Materials and Methods 43-54

4. Results and Discussion 55-91

4.1 Diversity of crabs, their identification and taxonomic revision.

4.2 Physico-chemical parameters in relation to abundance of

Maydelliathelphusa masoniana

4.3 Population structure of M. masoniana from natural habitat.

4.3.1 Sex ratio

4.3.2 Size frequency distribution

4.3.3 Breeding Season.

4.4 Studies on morphometric parameters viz., size, sexual maturity,

allometric growth and length–weight relationship of freshwater crab,

Maydelliathelphusa masoniana.

4.5 Evaluation of nutritional status viz., Protein, Lipid, Ash & Moisture

content in the edible meat of Maydelliathelphusa masoniana and its

relationship with spawning activity.

4.6 Macroscopic & Microscopic analysis of gonad development of

Maydelliathelphusa masoniana.

5. Summary and Conclusions 92-95

6. Recommendations 96-98

7. Bibliography 99-142

8. Appendix 143-154

Chapter 1

Introduction

Introduction

1

Life exists in very diverse form on earth showing huge biodiversity. Biodiversity

is a composite measure of the number of species, in terms of richness and number of

individuals in terms of relative abundance. Fresh water systems in tropics host a diverse

endemic fauna including fresh water crabs with 1280 species representing one fifth of all

the world’s brachyurans. (Cumberlidge, 2009). Of more than 6,700 known species of

brachyuran crabs, over 1,300 are true fresh water crabs. At global position fresh water

crabs account for a total of 238 genera, including 1,306 true fresh water crab species

accommodated in eight families. Out of these three families viz. Potamidae,

Gecarcinucidae and Parathelphusidae are found to inhabit Asia and Australia continent

(Yeo et al., 2008). From India, a total of 96 species under 41 genera in 6 families have

been recorded (Wood-Mason, 1871; Henderson, 1893; Alcock, 1910; Bott, 1970; Bahir

and Yeo, 2007 and Ng et al., 2011). The term fresh water crabs refer to those crabs that

have adapted fresh water, semi terrestrial or terrestrial modes of life and are characterized

Introduction

2

by their ability to complete their life cycle independent of the marine environment. The

colonization of fresh water has required crabs to alter their water balance by achieving

the ability to reabsorb salt from the urine, thus restricting water loss, though they need to

return to water periodically in order to excrete ammonia. On the other hand presence of

pseudolungs in addition to gills in most of fresh water species has developed to the extent

that it allows some species to be almost exclusively terrestrial (Cumberlidge, 1991).

Freshwater crabs belong to the order decapoda, the crustacean group that also includes

lobsters, prawns, crayfish and hermit crabs, which share the characteristic presence of

five pairs of thoracic legs (pereiopods). In freshwater crabs, the first pereiopods are

modified as pincers (chelipedes), and the remaining four pairs are relatively unspecialized

walking legs. Decapod crustaceans generally show sexual dimorphism in their external

morphology. The general body plan of fresh water crabs consist of a head, thorax and

abdomen , with the head and thorax (cephalothorax) covered by a broad carapace and the

abdomen reduced, flattened and flexed under the thoracic sternum. In adults, the male

abdomen is slim and narrow, and is either triangular or T shaped, while the female

abdomen is broad and round and covers nearly the entire thoracic sternum.

Decapod crustaceans represent a large number of living species that inhabit a

wide variety of biotopes (Mantellatto and Souse, 2000). Compared to marine crabs,

freshwater and terrestrial species show the following evolutionary trends, habitat

specialization, brood protection and abbreviated larval development (Hartnoll, 1988).

These decapods have been reported in almost all fresh water bodies from clear

fast flowing streams to sluggish lowland rivers and streams, fresh water swamps,

stagnant ponds and rice fields and even in pools in tree holes and leaf axils. They are

primarily, emerging to feed at night, omnivores and some species provide important food

sources for various vertebrates (Yeo et al., 2008). A number of fresh water crabs are

secondary hosts of flukes in the genus Paragonimus which causes paragonimiasis in

humans (Dobson, 2004.) Fresh water crabs constitute a very important group, both from

ecological as well as economical point of view. They play a significant role in nutrient

cycle, water quality monitoring and fishery wealth as they are consumed in many parts of

the world.

Introduction

3

Fresh water crabs though have remained a neglected component of the world’s

inland aquatic ecosystem, but because of their such characteristics as rapid growth,

production of large number of youngs, early attainment of sexual maturity, they qualify

themselves as culturable candidate.

The crustaceans are highly sensitive to pollution and hence any disturbances in

the physicochemical parameters in turn may induce changes in immune status of

crustaceans, by stressing them and resulting in a reduction of immune vigour. The

environmental parameters of any wetland are very important because the variations in the

physicochemical properties such as temperature, salinity, pH, dissolved oxygen, FCO2,

Ca++

, Mg++

, influence the crustacean abundance and life cycles. The correlation between

physicochemical parameters and crab population provides information regarding the

optimum conditions under which fresh water crabs can reproduce and thus maintain the

population.

Information on the population biology of fresh water crabs is very scarce

(Gherardi and Micheli, 1989). Studies on population generally focus on description of

density, size structure, sex ratio and breeding periods (Branco et al., 2002). Majority of

crab species are narrow endemics, occurring in only a small geographical area. This can

be attributed to their poor dispersal abilities, low fecundity and habitat destruction caused

by human population. Population dynamics of any species can be helpful in chalking out

the strategies to verify the factors accounting for the differences among population and to

understand the biology of constraints that are shaping the structure of these populations.

In crustaceans, as growth progresses, certain dimensions of the animals body may

grow much more than others, resulting in the phenomenon known as relative growth

(Hartnoll, 1974). Sexual difference observed in the growth of several body parts relative

to carapace size have often been used to examine the relationship between morphometric

and sexual activity in addition to morphometric difference among populations or species

(Aikens and Waddy, 1989). Studies of relative growth in crustaceans allow to define the

type of allometry in the growth of different body parts, such as chelae, locomotor

appendages, abdomen and pleopods, and to relate them to their specific functions. One

probable factor responsible for these changes in the allometric growth is the sexual

Introduction

4

maturity (Gonzalez-Gurriaran and Freire, 1994). In population studies, while

morphometric analysis provide a powerful complement to genetic and environmental

stock identification approaches (Cadrin, 2000) the length weight relationship allow the

conversion of growth in length equations to growth in weight for use in a stock

assessment model (Moutopolos and Stergiou, 2002).

The mathematical length-weight relationship yields information regarding the

general well-being of individuals, variation in growth according to sex, size at first

maturity, gonadal development and breeding season. Such studies in aquatic animals

have wide application in delineating the growth patterns during their developmental

pathways (Bagenal, 1978) and can be used to estimate the recovery of edible meat from

crabs of various sizes (Lagler, 1968). The length width /weight relationship is regarded

as more suitable for assessing not only fish but also crustacean (Sukumaran and

Neelakantan, 1997; Tabash, 2001). These relationships are often used to calculate the

standing stock biomass condition indices, analysis of ontogenic changes and several other

aspects of crustacean population dynamics (Atar and Secer, 2003).

Reproductive cycles of a crustacean include a series of morphological and

physiological events. This cycle is basic to all sexually reproducing crustaceans but the

time relationship between certain events and duration of these events varies for different

species (Sastry, 1983). Taking into account, the degree of gonad development, the

macroscopic characterization of the gonads among decapods have been commonly

investigated to determine the onset of the sexual physiological maturity or histological

maturity. (Costa and Negreiros-Fransozo, 1998; Castiglioni and Santos, 2001; Castiglioni

and Negreiros-Fransozo, 2006).

Crabs are the basic components of the ecosystem and they are consumed as food

in many countries. Edible crustaceans such as crabs, prawns, shrimps, crayfishes and

lobsters comprise major sources of nutritious food for humans. The nutritive value of

crustaceans depend on their biological constituents such as proteins, carbohydrates,

lipids, vitamins and minerals. Crabs have exceptional and scrumptious taste as compared

to fish and molluscs and rank third after shrimps and lobsters for their revered delicacy

and value of fishery they support (Savad and Raghavan, 2001). Crab meat contains many

Introduction

5

nutrients and is an excellent source of high quality proteins, vitamins and minerals. Yeo

et al., (2008) reported that a large number of potamids and parathelphusids are consumed

in Thailand and potamids are consumed by natives of South America to improve health

and cure physical injuries. Many curative properties are attributed to crab meat in view of

the fact that it is used to treat asthma and chronic fever (Raja, 1981). Crab fishery in India

is fast developing and there is immense scope for crab meat due to its delicacy and

nutritional richness. Because of their delicacy and large size, crabs are in great demand in

both National and International markets.

Knowledge of the biochemical composition of any edible organism is extremely

important since these reflect the nutritive value (Nagabhushanam and Mane, 1978). In

India, a total of 12 marine/brackish water species belonging to six genera, viz Scylla,

Portunus, Charybdis, Matuta, Varuna and podophthalmus have been extensively studied

for proximate composition of amino acids and cholesterol contents (Srinivasagam, 1979;

Sudhakar et al., 2009, 2011) but a very limited research, has been focused on freshwater

crabs (Sayyad et al., 2008; Sengul and Zeliha, 2011; Manhas, 2012).

In Jammu region of J&K State (North India), fresh water being the only source of

aquatic fauna, people mostly rely on fishes as food. Very scanty work has been reported

on fresh water crabs except for the presence of Paratelphusa masoniana Henderson

(1893) in Gadigarh stream (Dutta, 1978), in Gho-manhasan stream (Gupta, 2012), its

nutritional status (Manhas, 2012), report of Potamon species of fresh water crab from

Poonch stream (Anjum, 2012) and from Anji stream in Reasi (Kotwal, 2014).

Gho-manhasan stream (lotic) located at a distance of about 12 Km (30067

/ Lat N;

74079

/ Long E) fed by river Chenab is a sole source of water for the people inhabiting its

adjoining areas, for irrigation as well as domestic purposes. It observes large scale

anthropogenic activities at many sites. Maydelliathelphusa masoniana, a fresh water crab

from Jammu region of J&K (North India) has remained unexplored, barring a few reports

on its occurrence and abundance (Gupta, 2012) nutritional status (Manhas, 2012) and

parasitic infection (Anjum, 2012). A lacuna is clearly evident particularly with regard to

the various biological aspects related to the maturity and breeding of the species besides

reproductive biology. The population biology of these organisms is still poorly known

Introduction

6

and when we look at freshwater crab, Maydelliathelphusa masoniana, very negligible

information is on record from India, in general and Jammu region of J&K State, in

particular.

Present investigations therefore, have been carried out with a view to generate

information on different biological aspect of freshwater crabs keeping in view the

following objectives.

To prepare an inventory of the crab resources available in various water bodies of

the region.

To correlate the seasonal occurrence and abundance of crabs with the ambient

ecological parameters.

To study various morphometric parameters viz. size, sexual maturity, allometric

growth and length weight relationship of fresh water crab Maydelliathelphusa

masoniana.

To investigate the various relevant parameters of population structure viz. size

frequency distribution, sex ratio and breeding seasons of Maydelliathelphusa

masoniana from their natural habitat.

To evaluate the nutritive value of the edible meat of adult male and female

Maydelliathelphusa masoniana and to establish its relationship with spawning

activity.

To make seasonal studies on macroscopic and microscopic analysis of gonad

development of Maydelliathelphusa masoniana.

Chapter 2

Review of Literature


7

1.1 Diversity

Origin of freshwater crabs dates back to 65 million years, between the end of the

cretaceous and the beginning of Tertiary period (Bott, 1955). Pretzmann (1962) reported

two different families of freshwater crabs in Iran viz., Gecarcinucidae and Potamidae

whose ancestors probably originated from the sea independently. Many species of the

Potamidae family migrated to different geographically isolated regions where they

adapted themselves with prevailing conditions (Bott, 1970; Pretzmann, 1976). The

freshwater crabs in the Euro-Asian areas have been extended from the Mediterranean

region (Krupp et al., 1987) to East –Himalaya (Bott, 1966). Pretzmann (1973) considered

that the freshwater crab families have originated from a number of different marine

ancestors and that morphological similarities appear to be the result of convergence, and

not common ancestry.


8

The major bio-geographic division in the Himalayas is between eastern humid

tropical fauna, derived from Indo-Chinese Malaysian area and a western stepped fauna,

derived from Ethopian/ Mediterranean area (Mani, 1974). Geologically it appears that

some of the larger Himalayan rivers like the Indus, the Brahmaputra and the Ganges

existed before the final elevation of the Himalayan mountains (Das, 1990; Shroder,

1993).

The pseudothelphusoid and gecarcinucoid freshwater crabs found today in the

majority of Gaondwanan fragments (South America, Africa, Madagascar, India, South

East Asia and Australia) are postulated to have an Gondwanan origin with present day

distribution patterns resulting from the breakup of the supercontinent (Rodriguez, 1986;

Ng et al., 1995). Ng and Rodriguez (1995) indicated that the dominance of gecarcinucoid

crabs in the Indian peninsula and the absence of potamids could be explained on the basis

of their long isolation on Gondwanan continent fragment before it collided with

continental Asia, where potamid are found in large numbers.

In addition to dispersal, distribution limits of true freshwater crabs are also

influenced by host of other factors. These include abiotic factors such as climate,

hydrology, tropography and altitude as well as biotic factors such as habitat, vegetation

and inter-specific competition. (Barbaresi and Gherardi, 1997; Cumberlidge, 1999; Dai,

1999). All these factors have led to a high degree of endemism in freshwater crabs. One

of the key process driving freshwater crab diversification being allopatric speciation

resulting from geographic isolation resulting in relatively low fecundity and poor

dispersal abilities of freshwater crab (Cumberlidge et al., 1999). Sternberg (2001) and

Cumberlidge (2001) on the basis of morphological studies recognized eight families in

two main lineages (i) the monophyletic Trichodactylidae in the marine Superfamily

portunoidea. (ii) a monophyletic group consisting of all three superfamilies viz.

Potamoidea, Gecarcinucoidea (Parathelphusidae and Gecarcinucidae) and

Pseudothelphusidea (Pseudothelphusidae).

The general distribution pattern of the freshwater crabs correspond well to the

general bio-geographic model on the Himalayas (Brandis, 2000). The mediteranen fauna

in the Western Himalayas is represented by the subgenus Potamon occurring in Iran,


9

Afghanistan and the right tributaries of the Indus river (Brandis et al., 2000). Such a

tropical fauna is represented in the freshwater crab by the genus Potamiscus with

Potamon koolooensis occurring in the region of Shimla.

Brandis (2001) reported one endemic subgenus within the genus Potamon,

Potamon (Himalayapotamon) Pretzmann (1966) from the Himalayas. The subgenus

included two species Potamon atkinsonianum (Wood Mason, 1871) and Potamon

emphysetum (Alcock, 1909). The subgenus Himalayapotamon is raised to generic level.

Based mainly on the morphology of the male copulatory system Himalayapotamon

belongs to the family Potamidae and zoo-geographically Himalayapotamon appears to be

an endemic genus related to Potamon.

Yeo and Ng (2003) suggested the distributional pattern of Potamidae where one

sub family (Potaminae) occurring in North Africa, Southern Europe, Middle East and

Himalayas and other sub family the Potamiscinae in South East Asia, China & Japan.

Freshwater crabs are found in all major habitat types including flood plains, swamps,

lakes, moist forest rivers, highland and mountain systems, large lakes and large rivers,

rapids of tropical and subtropical areas. (Theme et al., 2005., Abell et al., 2008).

Brandis and Sharma (2005) while dealing with Maydelliathelphusa examined

many species from Nepal and Northern India and concluded that this species is

distributed along the Ganges and Brahmputra valleys ranging from Punjab to Nagaland.

It predominantly inhabits the lowland banks of rivers but reaches higher attributes (up to

3500m) in the Himalayas. Srivastava (2005) while conducting numerical status of species

distribution of freshwater crabs (Potamonids) in South India reported about 550

specimens comprising of 5 species belonging to 4 genera, under one family. The five

species being Travancoriana schirnerae, Balytelphusa guerini, B. cunicularis,

Oziotelphusa sensex sensex, Spiralothelphus hydrodroma.

Bahir and Yeo (2007) while studying the freshwater crab fauna of the two

southernmost Indian states of Kerala and Tamil Nadu reported nine genera and twenty

three species of gecarcinucid freshwater crabs, including six new genera and 10 new


10

species. Ng et al., (2008) elevated Maydelliathelphusa to the genus and listed five species

M.edentata, lugubris, falcidigitis, harpax and masoniana.

The oriental region is taxonomically most diverse part of the world with over 900

species of freshwater crabs in 154 genera and two families. In bio-geographical terms the

Indo-Burma hotspot lies in oriental region including India, Southern China and South

East Asia. The Indo-Burma 182 known species of freshwater crabs in 55 genera and two

families, the Gecaicinucidae and the Potamidac (Yeo, et al., 2008; Cumberlidge et al.,

2009) with 76% and 92% of endemism respectively have been recorded. According to

Yeo et al., (2008), Cumberlidge et al., (2009) and Klaus et al., (2009), south east asia

represents a biodiversity hot spot for extant primary freshwater brachyurans that lack any

marine members and occur as the families Potamidae and Gecarcinucidae comprising

about 12% of the total number of currently described brachyuran species.

Yeo et al., (2008) presented an assessment of the global freshwater crab diversity.

From all zoogeographical regions (except Antarctica), a total of 1,476 species in 14

families are currently known, including 1,306 species in eight exclusively freshwater

families (Pseudothelphusidae, Trichodactylidae, Potamonautidae, Trichodactylidae,

Potamonautidae, Deckeniidae, Platythelphusidae, Potamidae, Gecarcinudidae and

Parathelphusidae). Estimates of true freshwater crab diversity including undescribed taxa

suggest the field being in a “Discovery” phase.

Rahman et al., (2008) assessed the status of freshwater crab resources in some

wetland ecosystem of Bangladesh. The study on species diversity included its taxonomic

position, morphometric and meristic characteristics, habits and habitat, distribution and

economic importance. A total of five species of crabs were reported with three belonging

to family Potamidae and other two to family Grapsidae and Parathelphusidae.

Nasrollahzadeh et al., (2011) gave first report on freshwater crab in the altitudes

of Gulain (Lakan area). Macrobenthos sampled from the freshwater stream included

freshwater crabs. The crab species identified was Potamon bilobatum belonging to family

Potamidae. Klaus et al., (2011) reported the first evidence for Miocene brachyuran

freshwater crab from South East Asia, a hot spot of extant freshwater crab biodiversity.


11

The report confirmed the presence of potamoid freshwater crabs in South East Asia

during Miocene, as suggested by previous molecular clock estimates.

Ng and Lee (2012) described a new species of the freshwater crab of genus

Phricotelphusa. Alcock (1909) (Gecarcinucidae) from a limestone cave in Perlis,

northern Peninsular Malaysia. The new species was characterized by very long

ambulatory legs, and differs from congeners in the structure of its carapace, male

abdomen and gonopods.

Takeda and Shantikumar Singh (2013) reported some freshwater crabs from

North East India Maydelliathelphusa lugubris (Wood Mason, 1871) of the

Gecarcinucidae and Potamiscus manipurensis (Alcock, 1909) and Alcomon

superciliosum (Kemp, 1913) of the Potamidae were recorded from Manipur State, North

East India bordered on Myanmar Parathelphusa burmensis (Bott, 1966) of the Potamidae

from Myanmar bordered on India were recorded and their taxonomic problems were also

discussed.

Guner (2014) reported current distribution of Potamon species in Turkish Thrace

from 18 streams and 4 lakes, Potamon fluviatile being one of four freshwater crab species

found in the Balkan Peninsula. Another species P. ibericum was found in Marica River in

Bulgaria and in Meric River in Turkey.

1.2 Physico-Chemical Parameters

The distribution and existence of crabs depend on specific environmental

parameters such as salinity, pH, temperature and dissolved oxygen (Diaz and conde,

1989; Lee and Winckin, 1995; Carmona-Suarez and Conde, 1996). While hatching and

larval development occurs successfully under a relatively narrow range of salinity and

water temperatures (Lee and Winckin, 1995), juveniles or adult crabs can tolerate a wide

range of salinities and temperatures. The physico-chemical parameters being vital

ecological factors directly affect oxygen consumption, metabolism, growth, moulting

hormones and survival of crustaceans (Chen et al., 1995).


12

Gopinathan et al., (1978) reported direct relationship between temperature and

juveniles of brackish water prawn. The water temperature and air temperature in general

go more or less hand in hand (Singhal et al., 1986). Truscott and White (1990) while

studying the influence of metal and temperature stress on the immune system of crab

abundance observed marked fluctuation in circulating haemocyte numbers in control

crabs. A gradual rise in temperature from 10 to 200C significantly increased blood cell

numbers although rapid change initiated no response. Besha and Qureshi (1993) observed

that the temperature ranging between 24-280C was conducive for the abundance of

freshwater prawns.

Anger (1991) studied the effect of temperature on the larval development of the

Chinese mitten crab, Enocheir sinensis and reported that the development from hatching

to metamorphosis occurred only at temperature more than 120C. With increasing

temperature, both overall survival and range of salinity tolerance increased whereas

development decreased exponentially. Temperature being a limiting factor in the aquatic

environment, affect the metabolic activities, growth, oxygen consumption, reproduction,

moulting, survival, distribution and migratory behaviour of crustaceans (Haffner, 2000)

Jong and Arvey (2002) considered temperature as the dominant environmental

variable affecting metabolic rate in poikilotherms and they suggested that temperature

affects lipofuscion accumulation in the Blue Crab, Callinectes sapidus. Braide et al.,

(2004) stated that temperature range of 26.64±1.180C and 30.83± 1.47

0C being optimum

for shell fish propagation corresponding with the report of Ansa (2005) who gave

temperature range of 25.90C & 32.4

0C. Brylawski and Miller (2006) while studying

temperature dependent growth of the blue crab Callinectes sapidus indicated a moult

process model predicting crustacean growth, including the temperature dependence of

intermolt period that can produce the extended over wintering phenomena during which

growth ceases. Dibia (2006) and Jamabo (2008) also reported a temperature range

between 270C and 30

0C being ideal for growth and well being of shell fishes.

The pH being another important parameter affecting crab diversity and

distribution in an ecosystem . In decapods, pH influence the metabolism, physiology and

maturation process (Muthu, 1977). According to the EPA (1980), accepted water quality


13

criteria indicated a pH of <6.5 units to be harmful to many species of fish. Therefore, the

pH range of 6.5-9.0 units was found to be suitable for the protection of aquatic habitats.

The pH being the scale of intensity of acidity and alkalinity of water and a

measure of the concentration of hydrogen ions. Most of the biological processes and

biochemical reactions being pH dependent (Minns, 1989). The alkaline pH was found to

be associated with more number of crab species. However, with increasing pH, the

number of species has been reported to decrease (Das and Sahoo, 1997). According to

Chang (2008), the increasing pH appeared to be associated with increasing use of alkaline

detergents in residential areas and alkaline material from wastewater in industrial areas.

Abowei (2010) reported a pH higher than 7 but lower than 8.5 to be ideal for biological

productivity while pH lower than 4 to be detrimental to aquatic life. Most organisms

including crabs & shrimps do not tolerate wide variations of pH over time and if such

conditions persist death many occur.

Dissolved Oxygen (DO) concentration clearly influence the behaviour of

decapods and life strategies on basis of oxygen consumption and energy content. Oxygen

deficiency leads to stressed conditions in crustaceans (Pearson and Rosenberg, 1978).

Anonymous (1988) reported that according to classified continental inland water sources

of the water pollution control regulation, if dissolved oxygen is 8mg/l the water is I class,

if it is 6mg/l, the water is II class; if it is 3mg/l , the water is III class and if dissolved

oxygen is <3mg/l the water is IV class.

Oxygen consumption has been reported in many species of crustaceans

(Sumpton and Smith 1990). The respiration rates influence the metabolism in crustaceans

and anaerobic metabolism bringing reduction in growth & moulting frequency and

finally causing mortality (Allan and Maguire, 1991). According to Howell and Simpson

(1994) at about 3mg/l bottom fishes start to leave the area and the growth of sensitive

species such as crab larvae get reduced. At 2.5 mg/l, the larvae of less sensitive species of

crustaceans start to die and the growth of crab species get severely limited. Dissolved

oxygen is therefore one of the important parameter for water quality assessment

reflecting the biological and physiological processes prevailing in the water. In


14

freshwater ecosystems, the minimum dissolved oxygen should not be <5.0mg/l for

survival of aquatic life (Egemen and Sunlu, 1999).

Cheng et al., (2003) reported that dissolved oxygen affects haemolymph

osmolality, sodium concentration on acid base balance, glucose and lactose levels in the

tissue. In Haliotis diversicolor supertexta the stress of dissolved oxygen at 3.08 mg/l

and lower causes disturbance in acid base balance as well as anaerobic metabolism

resulting in glucose and lactate productions and acidosis in short term period. The value

of dissolved oxygen (DO) is found to be remarkable in determining the water quality

criteria of an aquatic system. In the system where the rates of respiration and organic

decomposition are high, the DO values usually remain lower than those of the system

where the rate of photosynthesis is high (Mishra et al., 2009).

According to Taylor (1949), total hardness being the parameter of water quality

used to decrease the effect of dissolved mineral (mostly Ca & Mg), determining

suitability of water for domestic, industrial and drinking purpose attributed to the

presence of bicarbonates, sulphates, chlorides and nitrates of calcium and magnesium.

According to Ryhanen (1962), calcium may limit the distribution and success of

crustaceans in soft water localities. This prevailing calcium deficiency in soft water

localities could cause delayed recovery of gastropods and crustaceans in acidified

freshwater. Incomplete calcification would cause prolonged periods of soft exoskeleton,

and make freshwater crustaceans more vulnerable to predation.

Colvocoresses et al., (1974) in blue crabs reported that the blood concentration of

magnesium decreases with an increasing calcium concentration in the external solution

and magnesium deposition may substitute for calcium deposition in the absence of

sufficient calcium. Smith et al., (1976) observed a noticeable softening of the

exoskeleton at low calcium levels of 5-7 mg/l in prawn, Macrobrachium.

Cameron (1985) while describing calcium uptake in the blue crab and other

crustaceans reported that the net uptake of calcium appeared to increase at highest

external concentration of calcium, but there was no increase in calcium concentration in

the blood, indicating that there must be an increased deposition onto the shell. The rate of


15

movement across the carapace epithelium into the shell would be expected to be

dependent on changes in the calcium concentration of the blood. The rate of calcification

is also limited by bicarbonate uptake apparent by H+ excretion.

Neufeld and Cameron (1994) studied the effect of the external concentration of

calcium in blue crab. Callinectes sapidus. Crabs transferred to water with 0.10m mol/l

calcium for the first two days after moult accumulated only 2.5g calcium /kg wet mass,

about one quarter of the mass accumulated. A period of acclimatization is necessary for a

component of the active transport system to increase its capacity.

Ali et al (2000) and Iqbal et al., (2004) stated that more than 15 mg/l CaCO3

hardness being suitable for fish and shell fish growth, and require liming for high fish

production. Rukke (2002) indicated that low calcium concentrations limit the distribution

and success of calcium demanding freshwater crustaceans in soft water localities.

Water with salinity below 1% are fresh and water with salinity higher than 1% are

brakish/marine. Ramane and Schlieper (1971) opined the salinity to be a major

environmental factor restricting the distribution of marine and aquatic taxa. Being an

ecological master factor in the distribution of living organisms, salinity, likely to

influence decapod crabs distribution with high levels during summer season and low

during the monsoon season (Chandra and Sreenivas, 1998). Chatterji et al., (2004)

studied the effect of salinity on larval growth of horse shoe crab, Tachypleus gigas and

reported that at 40% salinity, the growth rate was faster and molting occurred earlier.

According to Raj Kumar et al., (2004) high concentration of chloride is considered to be

the indicator of pollution due to organic wastes of animal or individual origin. Chlorides

being troublesome in irrigation water are also harmful to aquatic life.

Dobson et al., (2007) studied crab abundance of Potamonautes odhneri in rivers

draining Mt Kenya. They were recorded from 14 of the 21 sites including all forest sites

than in any agricultural site constituting about 70% and 40% respectively of total

macroinvertebrate biomass. Nayan et al., (2008) investigated the crab population in

relation to its ecological characterstics of Paratelphusa (Sartoriana spinigera) in Kawar

lake, India. It was observed that the total crab population exhibited positive correlation to


16

water temperature (r=0.121), conductivity (r=0.491) total hardness (r=0.202) and calcium

hardness (r=0.163), while negative correlation was exhibited with pH(r=-0.1242) and

chloride (r=-0.3612).

Lawal-Are and Kusemiji (2010) studied the effect of salinity on the survival and

growth of callinectes amnicola . The crabs were euryhaline and tolerated a salinity range

of 5 to 25% and had 90% survival at 15-20% of salinity. The highest gain in weight

(173.0%) and carapace width (56.1%) was obtained at salinity of 15% . Also complete

moulting was obtained at salinity of 15%. According to Lim and Wong (2010) the

temperature of 240C to 29

0C is optimum surface temperature to stimulate emergence of

Uca species from burrows. The pH of ground water was correlated with the crab

community structure. Most species were encountered in zone with high pH values. Low-

pH conditions were stated to be detrimental to crabs because they depend on calcium

carbonate for the formation of exoskeleton and shell building.

John et al., (2011) studied hydrological parameters at the nesting grounds of

horseshoe crabs along the Pahang coast, Malaysia. The mean annual temperature was

24.08± 2.910C with the mean annual salinity of 17.9± 10.63ppt. Seasonal fluctuations in

the pH of the water varied from 8.65-6.61. The mean annual dissolved oxygen (DO) in

the water varied from 6.24±0.95ml/L. Physico chemical parameters showed water

(P<0.05) while other parameters such as temperature, pH and dissolved oxygen did not

vary significantly.

Gupta et al., (2013) studied effect of physico-chemical parameters on crab

abundance of Paratelphusa masoniana. Crabs were observed to exhibit a negative

correlation with air (r=0.51465) and water temperature (r=-0.50615), depth (r=-0.62443),

pH (r= -0.33301) FCO2 (r=-18454) and Cl ( r=-0.1896) compared to DO, Ca2+

, Mg2+

&

HCO3 which were observed to bear a positive correlation (r=0.5072, 0.3393, 0.547395

and 0.030075 respectively). Sakhare and Kumble (2013) studied the influence of

physico-chemical parameters over reproduction of crab Barytelphusa cunicularis.

Physico chemical parameters as Temperature, pH, Dissolved oxygen, Free carbon

dioxide, Hardness, Chloride, Alkalinity, Nitrate and contamination of heavy metals like

Fe+, Na

+ & K

+ had direct or indirect impact over the metabolic content of targeted cells.


17

Altered physico-chemical parameter influenced gametogenic mechanism of crab B.

cunicularis showing hypertrophy of testicular cells thus leading to infertility.

Varadharajan et al., (2013) while studying the effect of physico-chemical

parameters on crabs biodiversity reported that unsuitable physico-chemical parameters

affect the distribution of crabs. He reported optimum minimum atmospheric temperature

(26.40C) and maximum (32.3

0C), salinity with minimum of ( 25.5%) and maximum of

(35.5%). The minimum pH was 7.6 and maximum of 8.3. Minimum dissolved oxygen

2.58 mg/l and maximum of 5.83 mg/l for the crab abundance.

Olatayo (2014) assessed the physico-chemical parameters of waters in Ilaje,

Nigeria to determine the levels of pollution through anthropogenic activities and state of

the aquatic ecosystem. Temperature recorded a mean of 29.750C with mean pH recorded

as 6.7. Salinity ranged from 16.35% to 16.65% thus indicating that all the physico-

chemical parameters showed no significant difference and the result obtained are within

the permissible level of aquatic biodiversity. Ngo-Massou et al., (2014) studied the

influence of some biotic and abiotic parameters on mangrove crabs. To verify the

interactions between crab populations and environmental components, the parameters

like salinity temperature, conductivity and pH were measured. Perisesarama kamermani

was found to be the most abundant (29%) dominating low salinity zone. Whereas,

portunus validus was present exclusively in high salinity zones.

1.3 Population-Structure

Studies on population structure of hermit crab have been conducted for tropical

(Kamlaveni, 1949; Ameyaw Akumfi, 1975) temperate (Asakura and Kikuchi, 1984),

European and Mediterranean (Lancaster, 1990; Elwood and Neil, 1991) crabs.

Populations of a given species at different latitudes subjected to particular environmental

condition may display different reproductive patterns. In most subtropical and tropical

regions, the reproductive activity is more intense during the warmer months, when the

food resource is abundant (Sastry, 1983). Information on the population biology of

freshwater crabs is very scare (Gherardi et al., 1987). Compared to marine crabs,

freshwater and terrestrial species show the following evolutionary trends: habitat


18

specialization limited geographic range, high rates of endemism, low fecundity with

small number of large eggs, brood protection and abbreviated larval development

(Hartnoll, 1988).

Turra and Leite (2000) explained that the occurrence of hermit crab populations

with seasonal reproductive patterns in the tropics and with continuous pattern in

temperate region are based on evolutionary histories and local factors such as competition

and shell use. Ample work has been reported on population and reproductive biology of

marine, interdial and estuarine crabs (Sallam, 2005; Henmi and Koga, 2009; Omolara,

2010) but limited published work is available on the population and reproductive biology

of freshwater, semi terrestrial or land crabs (Liu and Jeng, 2007; Wehrtmann et al.,

2010).

Sex ratio depicts the level of competition faced by crabs for reproduction

whereas, size frequency distribution helps in determining the dominant class size as well

as maximum size gained by both the sexes of crab. Sex ratio in majority of species is

close to unity, despite some variations between populations of a species and from year to

year in the same population. (Nikolski, 1963). According to Werner (1972), sex ratio

differing from the 1:1 are widespread among crustaceans. The overall sex ratio differed

from the excepted 1:1 proportion. Several causes may be held responsible for this

discrepancy, such as differences between sexes in longevity and growth rate, differential

migration, mortality and sex reversal.

According to Hicks (1985), the pattern of breeding activity is different in different

species of freshwater crab like T. Schirnrae. Some gecarcinucid crabs have short

breeding season during summer and some have long breeding seasons. Rao et al., (1986)

while studying reproductive cycle of the crab Ocypoda macrocera indicated breeding

period of this crab, to be a prolonged one extending from March–September with small

intermittent peaks and falls. Salinity was found to have a close relationship with the

breeding cycle as lower salinities October- January coincided with the absence of

reproductive activity.


19

Henmi and Kaneto (1989) reported that the presence of females with mature/spent

gonads and percentage of ovigerous females over the year is the most common technique

used to determine the reproductive period of a species. Jivoff (1997), while studying

sexual competition among male blue crab, Callinectes sapidus, advocated that in aquatic

brachyuran, males attained maturity at size bigger than females. Several authors have

extensively studied ovarian development to establish the reproductive period (Lopez-

Greco and Rodriguez, 1999).

Tongdee (2001) studied the size frequency distribution of mud crabs, Scylla spp.

and reported that percentage of smaller crab (smaller than 7cm CW) increased in May,

June and July followed by gradual decrease from August to October and again increase

during months of November. On the basis of these findings it was held that in these crabs

recruitment occurred twice in year, first during May to June and secondly in November.

Tongdee (2001) reported that mud crab Scylla species witness two recruitments in a year

indicating these mud crabs to be seasonal breeders, breeding twice in a year.

According to Mansur and Hebling (2002), low percentage of crabs in neotropical

freshwater of Brazil, during the breeding period may be attributed to the kind of

behaviour of these females as they remain in their burrows during the entire incubation

period and forage less. Sigana (2002) analyzed the abundance of crab, Thalamita crebata

in Kenya and observed that in smaller size i.e. 40.5-55.44 mm females were recorded to

be more numerous than males. Males, however, were reported by them to dominate in

larger size classes ranging from 55.4-80.44 mm.

Lardies (2004) studied sex ratio in Petrolisthes laevigatus and suggested that

deviation of sex ratio from 1:1 helps to internally regulate population by affecting its

reproductive potential. Litulo (2004) reported overall sex ratio of the fiddler crabs Uca

inversa to be 1:0.84 with further indication of male biased population. Studies on size

frequency distribution were carried out by Ali et al., (2004) on mud crabs, Scylla serrata

of Bangladesh . Based on their studies they reported that in males the maximum size

(130mm CW) was observed in the month of May and minimum (47 mm CW) in month

of August with the modal class size to be 81-90mm. In females the maximum size (100

mm CW) was recorded in month of June and minimum (32 mm CW) in July. Further, the


20

frequency of males was found by them to be more at class size 41-50 mm CW upto 81-90

mm CW and less thereafter, whereas in females the maximum frequency was recorded at

71-80 mm CW. Ali et al., (2004), while working on mud crab, Scylla serrata of

Sunderbans mangrove, reported that in these crabs, overall sex ratio of male with respect

to females was found to be 1:0.94 and was observed to fluctuate monthly.

Studies carried out on Dotilla Sulccata by Sallam (2005) revealed that overall

sex ratio which was recorded to be 1:1.1 remain fluctuated throughout the year. Also

numerically males were observed by them to be more in number than females except

during August and November where females surpassed them. Litulo and Tudge (2005)

studied the population dynamics of the diogenid hermit crab Diogenes brevirostris

focusing on size structure, sex ratio and breeding season. A total of 622 crabs were

obtained of which 290 were males (46.6%), 170 were non-ovigerous females (27.3%)

and 162 were ovigerous females. The overall sex ratio (1:1.15) differed from the

expected 1:1 ratio. Sexual dimorphism was evidenced by the larger size attained by males

in relation to both ovigerous and non-ovigerous females. Breeding occurred year-round

with three peaks of spawning (March, August and December).

Czerniejewski and Wawrzyniak (2006) while studying the seasonal changes in

the population structure of the Chinese mitten crab, Eriocheir sinensis exhibited that the

overall sex ratio differed from the expected 1:1 ratio, but not significantly and overall

occurrence of males were higher than females in autumn months.

Lawal-Are (2010) studied sex ratio in blue crab, Callinectes arnicola in Nigeria

and found it to be 1:0.96 exhibiting fluctuating pattern. Females were reported to be

significantly more abundant than males in dry months of March , May and August and

early month of rainy season in September. Venancio and Leme (2010) analyzed the

biology of freshwater crab Trichodactylus petropolitanus from a population inhabiting a

small montane stream in the Atlantic Forest of Brazil. The analysis of relative growth

showed that females reach morphological maturity at a smaller size than males. Size

frequency distributions indicated that females grew faster than males.


21

Pathre and Patil (2010) investigated the breeding cycle in freshwater crab

Barytelphusa cunicularis. The high percentage of mature female and male crabs was seen

during June, July and August indicating the reproductive period of this species.

Maximum number of ovigerous females were observed during July to September and the

breeding peak was correlated with maximum development of gonads. The freshwater

crab, Barytelphusa cunicularis was found to be continuous breeder and reproducing

females and males were found all the year around. Bessa et al., (2010) studied the

temporal and spatial variability in population dynamics of European Crab Carcinus

maenas. Juveniles recruitment was continuous throughout the year with a high proportion

of young recruits in the spring. The C. maenas population showed a regular size-

frequency distribution and structure for both sexes.

Stauffer et al., (2011) investigated the population structure, fecundity and relative

growth of Leurocyclus tuberculosus from coast of Rio. De Janeiro, Brazil. The sample

totalized 269 crabs , 168 males and 101 females (42 ovigerous). The sex ratio was 1:1.66

(M: F) and differed statistically from the expected (x2 = 16.68; P>0.05). The carapace

width (CW) ranged from 12.81 to 71.67 mm (mean ±SD 48.77± 13.75 mm) and from

15.33 to 55.44 mm (36.19± 8.66mm) in males and females, respectively. The size at

onset of sexual maturity was estimated at 30 mm (females) and 55 mm (males). Sharma

and Gupta (2013) while studying population structure of freshwater crab Paratelphusa

masoniana evaluated a total of 581 crabs of which 283 (48.37%) were males and 302

(51.62%) were females. The overall sex ratio was found to be 1:1.07 indicating that

females were numerically more as compared to males. However, sex ratio never remain

constant but exhibited fluctuation pattern throughout year especially in the month of

June-July and December-January when it was less than unity (1:0.40, 1,1:0.6 and 1:0.55

respectively). The present population exhibited non normal size frequency distribution

with male reaching greater size (5-6 CW) than females (4-5 cm CW). Devi and Smija

(2013) studied reproductive biology of freshwater crab Travancoriana schirnerae with

respect to breeding period and fecundity. A total of 3605 individuals were collected of

which 2168 (60%) were males and 1437 (40%) females. The breeding season extended

from February-May and the highest frequency of ovigerous females was observed in

March juvenile carrying females appeared in the population from April-July, with the


22

highest percentage in May. Recruitment of juveniles in the population was recorded at the

onset of rainy season in June.

Alarcon et al., (2014) studied the basic parameters of the population structure of

Trichodactylus fluviatilis in Ubatuba Brazil. A total of 306 individuals were captured

with 138 males and 168 females. Females mean size (23.53 ± 4.8 mm CW) was

significantly larger than males (22.32 ± 4.8 mm CW). There was no significant deviation

from a 1:1 sex ratio during the study period and the size distribution of the sampled

population presented a normal distribution.

1.4 Morphometric parameters

Information about individual body weight, length/width relationships in

population is important for estimating the population size of a stock, specifically such

information regarding the distinguishing characters and size relationships in sexually

mature individuals is of significant importance in commercially valuable crustaceans.

Such knowledge help establishing Taxonomy of a species, its life history patterns and

also serve important tool in fishery resource management and culture.

According to Huxley (1932), the allometric equation is the most utilized method

for analysis of growth during the ontogeny. To estimate the coefficients of allometric

equation, the data are usually logarithmized and subjected to linear regression (Teissier,

1960). According to Lagler (1968), the carapace and the length weight relationship can

be used to estimate the recovery of edible meat from crabs of various sizes.

Decapod crustaceans generally show sexual dimorphism in their external

morphology. Sexual difference observed in the growth of several body parts relative to

carapace size have often been used to examine the relationship between morphometrics

and sexual activity in addition to morphometric difference among population or species

(Kanno, 1972). In decapods, Hartnoll (1974) reported the importance of chelar

dimensions to characterize sexual dimorphism or to estimate size at sexual maturity.

Decapod species develop these chelipeds for combat, display and courtship.


23

The mathematical length weight relationship yields information on the general

well being of individuals, variation in growth according to sex, size at first maturity,

gonadal development and breeding season. Study of length-weight relationship in aquatic

animals has wide application in delineating the growth patterns during their

developmental pathways (Bagenal, 1978). Peters (1983) and Schmidt-Nielsen (1984)

described allometric regression changes in soft tissue content or total animal weight for

crustaceans relative to carapace width /length.

Shine (1988) proposed a pattern regarding the difference in the size at

morphological sexual maturity between females and males, according to which the

morphological difference between the sexes being the requirement for reproduction.

When females allocate their energy for reproductive purpose, they tend to mature at

smaller sizes than males, who invest their resources in somatic growth and reach maturity

at greater sizes. The length width/weight relationships are regarded as more suitable for

evaluating crustacean populations (Prasad and Neelakantan, 1988). The transitional

phase in Brachyura involves morphological changes that can be detected by inflections or

discontinuities in a series of linear or curvilinear relationships using a bivariate analysis

(Haefner Jr., 1990).

Positive allometric growth is an indication of a crabs heaviness with the

implication that the crabs are heavier than their lengths. The change in growth coefficient

(b) value depends primarily on the shape and fatness of the species, seasons or time of the

year, temperature, salinity food (quantity, quality and size), sex and stage of maturity

(Sparre, 1992). Studies of relative growth in crustaceans allow to define the type of

allometry in the growth of different body parts such as chelae, locomotor appendages,

abdomen and pleopods and to relate them to their specific functions. One responsible

factor for these changes in allometric growth is the sexual maturity (Gonazalez-

Gurriaran and Freire, 1994).

Allometric analysis contributes to a better understanding of many biological

events that occur in life cycle of animals (Gould, 1996). Body weight, total length and

carapace length are the most frequently used dimensions in the study of Crustaceans

(Sukumaran and Neelakantan, 1997). Chu (1999), analyzed morphometric character and


24

organ indices in Charybdia affinis collected from Zhuyiang estuary, China. There was

marked sexual difference in relative growth of the cheliped. The female abdomen

increased in size at puberty, accompanied by a reduced allometric growth rate. Analysis

of gonad development gave the estimate of 50% gonadal maturity at 42 and 36mm

carapace width for females and males, respectively.

In a study of the crab Chaceon affinis, by Fernandez-vergaz et al., (2000), it was

suggested that although the functional capacity of mating is related with the

morphological maturity, this is attained before the individual becomes physiologically

mature and is therefore able to reproduce. Rosenberg (2002) used modern methods of

describing shape and size, geometric morphometry to study claw variation across the

genus Uca. Within the species, major claws showed allometric trends in both shape and

size being isometric with respect to size , but allometric with respect to shape.

According to Atar and Secer (2003), the relationships between carapace length

and weight of the crabs have many uses. They are often used to calculate the standing

stock biomass, condition indices, analysis of ontogenic changes and several other aspects

of crustacean population dynamics including management of population. Carapace width

is the body dimension most used as the independent variable in the analysis of relative

growth of crabs, because it fully represents the physiological changes that occur

throughout their life history (Castiglioni and Negreires-Fransozo, 2004).

In crustaceans, the allometric relationship between body size and various organs

are used to estimate the sexual maturity, assuming that the secondary sexual

characteristics appear and grow at different rates in mature and immature stages (Leme,

2005). Ashkenazi et al., (2005) proposed for the first time a series of 22 easily

measurable morphometric parameters that permit assessment of the pincer size of fossil

fresh water crabs from measurable fragmented pincers. The pincer length represented the

crab body size in Potamon potamios. Regression line equations were given as an

implicative tool for future assessment of the pincer size of fragmentary fossil crabs from

the northern Jordan Valley.


25

Aiken-Oriola et al., (2005) carried out morphometric and meristic studies in two

crabs: Cardiosoma armatum and Callinectes pallidus. Maximum carapace length was 6.0

cm (4.38±0.04cm) in Cardiosoma armatum and 7.0 (5.17±0.04 cm) in Callinectes

pallidus. Mean body weights were 110.21±4.14g and 126.27±4.01g respectively. Body

weight, chelae diameter and condition factor were significantly higher in males of

C.pallidus. In the mixed population, length weight relationships indicated positive

allometric growth (b<3) in C.armatum and a positive isometric growth pattern in

C.pallidus (b=3).

The allometric relationships are powerful tools used by taxonomists and

ecologists in the analysis of intraspecific and interspecific variation among different

populations and to estimate the average size at sexual maturity related to environmental

conditions (Costa and Soares-Gomes, 2008). Olusoji, et al., (2009) studied length weight

relationships of the West-African freshwater crab, Sudanonautes africanus. Growth

parameters a and b of the length weight relationship was 0.889, 5.029 and 0713 for

males, females and entire population respectively. There was strong relationship

(r2=0.81) between length and weight of males and females and the entire crab population.

The length weight relationship (LWR) was allometric for all crabs.

Josileen (2011) studied the interrelationships between various morphometric

characters viz., carapace width and length and chelar propodus length and height in

males, as well as carapace width and length and abdominal width and length in females

were estimated using a total of 980 crabs, Portunus pelagicus. The allometric

relationships suggested that most relationships are positive and highly significant. The „b‟

values for carapace width-weight in males and females were 3.607 and 3.293,

respectively and for carapace length-weight they were 3.049 and 2.774, respectively. The

result showed a significant deviation from an isometric growth pattern, indicating a

significant difference between sexes with respect to the carapace width-weight

relationship.

Patil and Patil (2012) studied length weight relationship and condition factor of

freshwater crab Barytelphusa gurini. The exponent „b‟ value for males 2.83 and for

females it was 2.03 and combined crabs was 2.35. Growth generally showed negative


26

allometry in all crabs. The regression revealed high correlation and the coefficient of

determination (r2) very close to 1.

Pathre et al., (2013) carried out studies on morphometery of freshwater crab,

Barytelphusa cunicularis. In males all the segments in major and minor chela showed

positive allometric growth while in female, merus and propodus only showed positive

allometric growth. Analysis of covariance of growth indicated there was a significant

difference in growth pattern whereas in females the growth constants did not differ

significantly with respect to chela segments in both the sexes. Ribeiro et al., (2013)

studied size at sexual maturity and the allometric growth of the semi-terrestrial crab

Sesarma rectum. A total of 492 crabs, being 262 males and 230 females were obtained.

The specimens were measured at carapace width (CW), the left and right propodus length

and height (RPL, RPH, LPL and LPH), and abdomen width (AW) of females. In males,

based on the relationship between CW and length of right propodus (LRP), the

morphological size at the onset of maturity was 27.14mm. In females, the size at sexual

maturity was 22.97 mm, based on the relationship CW and AW.

Herrera et al., (2013) determined the size at sexual maturity in the freshwater crab

Dilocarcinus pagei. The dimensions measured were carapace width (CW), carapace

length (CL) propodus length (PL), and abdomen width (AW). The morphological

maturity was estimated based on the analysis of relative growth based on the allometric

equations Y=axb. Based on the relationships, the estimated value to morphological sexual

maturity was 21.5 mm (CW) in males and 19.7mm (CW) in females. The minimum size

for capture in D. pagei was reported to be 40mm (CW) based on the data obtained for

sexual maturity.

Mady-Goma et al., (2014) carried out morphometric study to describe freshwater

crab Sudanonautes aubryi. 186 specimens were identified and measured with sex ratio

equal to 1.62, and the average weight to be 6.3g. The average weight of females (8.13g)

is higher than that of males (5.03g). The average width of the carapace is 25.73mm; the

carapace of the female being larger (30.18mm) than males (22.99mm). Okon and Sikoki

(2014) studied length weight relationship and condition factor of the West African fiddler

crab (Ucatangeri). Ucatangeri exhibited sexual dimorphism with sex ratio of 1.2:1.0


27

which was significantly biased in favour of males. The length-weight relationship showed

negative allometric growth, b=1.6431.

1.5 Nutritional Status

The knowledge of the biochemical composition of any edible organism is

extremely important since the nutritive value is reflected in biochemical contents as

stated by Nagabushanam and Mane (1978). Crab meat is an excellent source of nutrition

because of its high protein content comparable with that of other sea-foods

(Srinavasagam, 1979).

Knowledge of the bio-chemical and metabolic processes during reproductive

cycle are essential for a complete understanding of crustacean reproduction. Many of

such studies concerned the accumulation of organic reserve constituents like lipids

followed by their metabolism and mobilization as one of the most significant metabolic

events in the physiology of crustaceans (Teshima et al.,. 1989). Biochemical changes

during maturation, molting and reproduction in gonads, hepatopancreas (HP) and muscles

have therefore been thoroughly worked out for a number of crustacean species (Castile

and Lawrence, 1989., Jeckel et al., 1991). Many investigators have estimated the

concentration of various elements in aquatic and biological samples collected from

various natural environments (Bu-olayan and Subramanyam, 1996; Hota et al.,. 2001;

Alasalvar et al., 2002; Ashok et al., 2003).

Proximate composition, amino acid and cholesterol contents have been

extensively studied in marine crabs from India (Srinivasagam, 1979; Sudhakar et al.,

2009; Sudhakar et al., 2011) and from abroad (Kucukagulmez et al.,. 2006 ; Dima et al.,

2009, Jimmy and Arazu , 2012). Limited research on freshwater crabs, however have

attracted the attention of workers in recent past only (Sayyad et al., 2008; Sengul and

Zeliha, 2011; Pati et al., 2012; Devi and Smija, 2013; Langer et al., 2013; Varadharajan

and Soundarapandian, 2014).

Systematic investigations on the biochemical composition of gonads in relation to

reproductive cycle has been initiated by Giese and his co-workers (Giese, 1959; Giese

et al., 1964; Pearse and Giese, 1966). Studies on the biochemical changes in relation to


28

reproductive cycles in invertebrates have been investigated by some workers. Rahman

(1967) studied the reproductive and nutritional cycles of the crab, Portunus pelagicus.

Similar studies were also carried out on another marine crab Charybdis variegata by

Chandran (1968).

Pillay and Nair (1973) studied the fluctuations in biochemical constitutes viz

water, protein, lipid and glycogen in gonads, muscles and hepatopancreas of Uca

annulipes , Portunus pelagicus and Metapenaeus affinis. The water and ash contents of

the entire body showed an systematic fluctuation in relation to the annual reproductive

cycle. During breeding season, there was mobilization of the lipid from hepatopancreas to

gonad to support intense gonad maturation. The maturing ovary contains more lipid than

an immature or spent ovary. The changes in the biochemical constituents in the testis

were not so pronounced as in the ovary, since the testicular cycle being almost

continuous in these crustaceans.

Diwan and Nagabhushnam (1974) investigated monthly changes in the

biochemical components of gonads of the fresh water crab, Barytelphusa cunicularis, to

determine their variability in the course of the reproductive cycle. During the ripening of

the gonads, glycogen, fat and protein contents were high. During spawning period all

values fell considerably reaching the minimum values at the end of spawning season.

During the maturation of the gonads, glycogen, fat and protein contents were increased.

According to Gibson and Barker (1979), the variations of hepato somatic index

throughout the year and during the maturation process, did not seem to corroborate the

general pattern among decapods, which was the storage of organic reserves in the

hepatopancreas and utilization of these reserves in ovarian development.

Akbar et al., (1988) investigated seasonal changes in biochemical composition viz

water protein, total nitrogen, glycogen, lipid and ash in both male and female crab of

Protunus pelagicus. The water content of the entire body showed no seasonal

fluctuations. Significant seasonal variations however were noted in other components of

edible parts viz protein (54 to 75%), glycogen (1.53 to 2.44%), total lipid (3.8 to 4.7%)

and total inorganic contents (8.4 to 11.3%).


29

Rosa and Nunes (2003) reported biochemical changes during the reproductive

cycle of Aristeus antennatus. Proximate chemical composition, lipid classes, fatty acid

profiles, glycogen and cholesterol content were determined in the muscle, ovary and

hepatopancreas. Both ovarian and hepatopancreas cholesterol increased with maturation

with major fatty acids 16:0, 18:1, 20:5 and 22:6, indicating these compounds to be the

major source of energy during embryonic and early larval development stages.

Sudhakar et al., (2009) investigated biochemical composition (protein,

carbohydrate and lipid) including amino acids and minerals in both soft and hard shell

crabs of Portunus sanguinolentus. The protein, carbohydrate and lipid contents were

found to be higher in hard shell crabs than that of soft shell crabs. Hard shell crabs had

maximum amount of essential amino acids (51.096%) as compared to soft shell crabs

(43.627%). Hard shell crabs contributed 53.783% of non essential amino acids in

comparison to 49.719% in soft shell crab.

Soundarapandian et al., (2010) studied effect of diets on the biochemical changes

of commercially important crab Portunus sanguinolentus. The protein content of the crab

offered with combined feed was maximum (41.08%) and poultry waste fed animals was

minimum (37.29%). The lipid content of the crab was similar for all feeds. No significant

difference was noticed with reference to ash content of the crabs.

Sengul and Zeliha (2011) investigated nutritional properties of fresh water crab

Potamon potamios in relation to season and sex. According to proximate composition

analysis, these were no significant difference in the moisture, proteins, fat and ash

contents in terms of seasons. But there were significant (P<0.05) changes between male

and female crab meats in term of protein values, and fresh water crab meat was found to

be a good protein source.

Ansari (2012) investigated seasonal variations in the protein content of freshwater

crab, Barytelphusa cunicularis and Barytelphusa guerinii. The protein contents of the

gonads of both the species showed a remarkable increase during the reproduction period.

The testis of Barytelphusa guerinnii had high protein content than ovaries. The ovary

protein content of Barytelphusa cunicularis was however, more than the testis during the


30

reproduction period. The results were discussed in the light of metabolism of the organic

reserves during reproduction. Pati et al., (2012) carried out biochemical composition of

fresh water crab meat Sartoriana spinigera (Wood Mason, 1871) before and after their

spawning seasons during the months of January to March and July to September. The

protein content of meat of female crabs varied from 30.03% to 59.29%; lipid 7.01% to

11.29%; ash between 38.75% and 39.44% dry weight and the moisture ranged between

71.26 % and 79.51%. The protein, lipid and ash contents were recorded highest during

the month of January to March whereas lowest percentage were observed during July to

September. The biochemical composition of crab meat having high protein justified the

edible quality of these crabs by humans.

Devi and Smija (2013) determined the nutritional status of claw and body meat in

adult male and female of the freshwater crab, Tranancoriana schirnerae with respect to

the proximate composition, free amino acids (FAA) and cholesterol content. The mean

protein, moisture, lipid, FAA and cholesterol contents of crab meat were found to be

19.39%, 83.02% 369.5mg, 1469mg and 21.3mg/100g respectively. Further, female crab

meat had significantly higher levels of protein, carbohydrate and FAA than males

indicating females being better than males nutritionally. Schirnerae meat being low in fat

, yet high in protein serve an excellent option to include in a healthy diet. Langer et al.,

(2013) investigated the seasonal fluctuation in the proximate composition of freshwater

crab, Paratelphusa masoniana. Marked seasonal variation in protein, lipid and moisture

were observed to determine their mobalization during the course of the reproductive

cycle. Maximum protein (62.15±0.30%; 55.85±0.48) and lipid (5.85±0.46%;

5.49±0.38%) were observed during non-spawning period and minimum during spawning

months. The relationship between protein and lipid was direct one but inversely related to

moisture content.

Omatayo et al., (2013) investigated proximate composition and mineral of the

land crab Sudanonautes africanus from exoskeleton, flesh and whole body. The results

showed that in the exoskeleton samples, crude protein was highest in the male with the

value of 39.84±0.25%. While in the flesh samples, carbohydrate was the highest in the

male with 41.59±0.27%. S.africanus was found to be rich in sodium, potassium, calcium,


31

magnesium and phosphorus. Information on nutrient composition of S.africanus

encouraged the processing, utilization and marketing of this commonest species of West

Africa. Nutritive values recorded for S. africanus suggested that the species could be

employed as an alternative dietary supplement of protein and minerals.

Varadharajan and Soundarapandian (2014) evaluated the nutritional status of the

edible potamid crab Spiralothelphusa hydrodroma from Prangipettai, South East coast of

India. The result showed that the constituent like proteins, carbohydrate, lipid, moisture,

ash and minerals were maximum in cephalothorax and minimum in swimming and

walking legs.

Proteins

Proteins being vital for normal function, growth and maintenance of body tissue.

Camien et al., (1951) reported that FAA content to be generally greater in marine species

than fresh water species. The nutritive value of animal meat being decided by its free

amino acid content (FAA). Crustacean muscles contain high concentrates of free amino

acids such as arginine, glycine, proline, glutamine and alanine (Cobb et al., 1975).

Konosu and Yamaguchi (1982) pointed out that the FAA content in crustaceans are

higher than those in fish and mollusc. Free amino acids such as alanine, glutamine and

glycine are responsible for the unique flavour and taste of crab meat. The content of

protein being considered as an important mean for assessing the physiological standard of

an organism (Diana, 1982). During the period of starvation, the abdominal muscles make

the largest contribution of protein to energy metabolism sufficient to make a substantial

contribution to the overall animal maintenance. (Barclay et al., 1983).

George and Gopakumar (1987) observed the protein content in Scylla serrata

with egg (19.16%), without egg (20.92%), body meat (16.8%) and claw meat (16.28%).

The protein content of Protunus pelagicus and Protunus sanguinolentus was 0.47 to

15.91% and 12.81 to 13.6% respectively. Prasad and Neelakantan (1989) reported that in

S. serrata, the protein content of body meat and claw meat was 20.11% and 18.54%

respectively with higher proportion of essential amino acids viz Histidine, leucine,

thronine & Cystine making a total contribution of 36.82%.


32

George et al., (1990) noticed the protein values in cooked crab of S. serrata in the

range of 14.43 to 18.96%. Khan (1992) investigated 11.60% protein in body meat of

male and 19.92% protein in female body meat of S. serrata. Anonymous (1997) reported

that the protein value in blue crab was 17.17%. Annon (1999) reported arginine, lysine,

leucine and isoleucine in blue crab.

Tureli et al., (2000) reported that breast meat protein content of Protunus

pelagicus (sand crab) were 18.83±0.23 and 17.55±0.23% in male and female

respectively. Vasconcelos and Braz (2001) reported the protein content in C. affinins to

be 17.8%. Balasubramanian and Suseelan (2001) while investigating S. Smithii assessed

the protein values to vary from 59.8 to 71% on dry matter basis.

Zafar et al., (2004) reported the protein values in S.serrata male to be 17.69% and

19.39% for females. Thirunavakkarasu (2005) recorded the protein values in S.

tranquebarica from different parts viz 65.48 to 72.24% for body meat, 69.5 to 80.29%

for claw meat and 69.47 to 74.7% for leg meat.

Shell fish meat provides high quality protein with all the dietary essential amino

acids for the growth and maintenance of the body (FNB, 2007). Kuley et al., (2008)

found differences between protein values of claw and body meat with higher value for

claw than body meat. Ozogul et al., (2010) reported that there were variations in protein

contents of both male and female crab meat in Callinectus sapidus. According to Sengul

and Zeliha (2011), the protein values recorded for female crab meat of Potamon potamios

were low in comparison to males of same species. Siva Sanker and Yogamoorthi (2012)

in the ghost crab Ocypode platytarsis revealed that lower FAA concentrations were found

in females than males.

Lipids

In crustaceans, lipids are not only the main organic reserve and source of

metabolic energy but also indispensable in maintaining cellular integrity. Lipids as a

general rule act as major food reserve along with protein and are subject to periodic

fluctuation influenced by environmental variable like temperature. (Jonston, 1917).


33

Lipids supply a source of essential nutrient particularly the fat soluble vitamins like A, D,

E and K (New, 1986).

George and Gopakumar (1987) assessed the lipid values in S. serrata with egg

(0.43%), without egg (0.7%), body meat (1.07%) and claw meat (1.0%). Prasad and

Neelakantan (1989) noticed that the lipid content in S. serrata in body meat was 1.65%

and claw meat was 2.01%. Annon (1999) reported that lipid value in blue crab was 1.5%.

Tureli et al., (2000) while investigating the fat content in the breast meat of blue crab

(Callinactus sapidus) and sand crabs (Portunus pelagicus) assessed the lipid value to be

1.51±0.01% and 1.53±0.01% respectively.

Lipids being highly efficient as source of energy, contain more than twice the

energy of Carbohydrates and proteins (Okuzumi and Fuji 2000). Balasubramanian and

Suseelan (2001) recorded the lipid values from 6.2 to 7.6% in Charybdis smithii. In

chaceon affinis the lipid values were 0.7% (Vasconcelos and Braz, 2001). According to

Adeyeye (2002) fresh water crab meat is low in fat, which is good for health.

Thirunavukkarasu (2005) estimated the lipid content of body meat (0.9 to 1.6), claw meat

(1.83 to 2.06%) and leg meat (1.58 to 2.08%) in S. tranquebarica but showed no notable

variation with sex. Murugesan et al., (2008) reported lipid content of hard shell crabs of

C. Lucifer in the range of 1:65%. The muscles of crab and prawns contained lesser

quantity of lipids (Bhavan, 2009). Fresh water crab meat being low in cholesterol when

compared to marine and brackish water species (Sinha and Ahmed, 2011).

Moisture

Radhakrishan and Natarajan (1979) observed comparatively lower moisture

values of 69.54 to 74.46% in P. vigil. Radhakrishnan (1979) also reported moisture level

of 69.52 to 80.51% and 67.44% to 82.04% in P.pelagicus and P. sanguinolentus

respectively. George and Gopakumar (1987) assessed the moisture values in S. serrata

with egg, without egg, body meat and claw meat as 78.02, 77.20, 80.19 and 82.94%,

respectively. Investigation on the edible crab, Portunus pelagicus by Akbar et al., (1988)

revealed that moisture content varied from 66.06 to 80.04% and from 66.40 to 78.55% in

all the three tissues viz. Body meat, claw, claw meat and trash of both male and female


34

crabs respectively. In S. serrata, Prasad and Neelakantan (1989) observed moisture of

77.10% in the body meat and 78.76% in the claw meat of blue swimming crab.

Balasubramanian and Suseelan (2001) while studying the biochemical

composition of the deep water crab, Charybdis smithii arrived at the conclusion that

moisture content ranged between 85.5 and 89.60% in case of male crabs in the intermoult

stage. Skonberg and Perkins (2002) found 79.0±0.7% moisture in Carcinus meanus.

Zafar et al., (2004) in S.serrata noticed higher percentage of moisture in body

meat than claw meat and observed negative correlation between water content and

protein concentration. Whereas, the moisture content of S. tranquebarica ranged from

73.5 to 81.8% in body meat, 73.5 to 80.16% in claw meat and 73.23 to 79.6% in leg meat

(Thirunavukkarasu, 2005). Kuley et al., (2008) reported that male blue crab meat had

lower moisture content than female crab meat. However, Bhavan et al., (2010) reported

that the muscles of male prawn generally contained higher levels of water then females.

According to Bassey et al., (2011) knowledge of the moisture content of food stuff serves

as a useful index of their keeping qualities and susceptibility to fungal infection.

Ash

Farragut (1965); Badawi and Alexendria (1971); Hamsa, (1978) while

investigating Protunus pelagicus and Callinectes sapidus respectively reported higher

values of ash for non-edible components. Ash is left out after complete combustion of

crab/fish meat and gives a measure of the total mineral content. The fish and shellfish can

absorb minerals directly from the aquatic environment through gills and body surface

(Gopakumar, 1997).

Investigations on the seasonal variation in biochemical composition of edible

crab, Portunus pelagicus Linneaus by Akbar et al., (1988) revealed that ash or total

inorganic content in Portunus pelagicus varied on an average from 8.44 to 33.29% (dry

wt.) in both male and female crab; higher values being recorded for non-edible

components.


35

Skonberg and Perkins (2002) found that ash content of S.spinigera varied between

38.75% and 39.44% (mean : 39.12±0.34%) with maximum as 39.44% in the month of

February and minimum of 38.75% in the month of January. According to Gokoglu and

Yerlikaya (2003) proximate ash contents of blue crab (Callinectus sapidus) and swim

crab (Portunus pelagicus) for claw meat and body meat were 1.39±0.019% and

1.89±0.08% respectively. Benjakul and Sutthipan (2009) reported that soft shelled crab

meat contained 1.67±0.9% of ash.

The ash content of S.spinigera varied between 22.01- 25.55% (mean: 22.27

±3.15%) with a maximum of 25.55% in the month of July and minimum of 19.25% in the

month of August (Pati and Routray, 2012). The ash content of S.africanus ranged

between 4.55±0.01% in male exoskeleton to 14.92±0.03% in the male whole body

(Omotayo and Abayomi, 2013). Varadharajan and Sondarapandian (2014) reported ash

content in S.hydrodroma to be maximum in cephalothorax (0.72±23.19) and minimum in

swimming and walking legs (0.31±0.29).

1.6 Gonadal development

Reproductive cycles of a crustacean comprises of a series of morphological and

physiological events. These events include proliferation of gonial cells (activation of

gametogenesis), differentiation and growth of gametes to maturation (gamete

production), reproductive behavior associated with mating, release of gametes

(ovulation), spawning, and incubation of embroyos until hatching to release larvae or

juveniles (Giese and Pearse, 1974; Sastry, 1983).

Factor such as temperature, salinity, food availability, photoperiod and lunar

cycles could determine the periodicity and extension of the reproductive period of a

species, as well as its fecundity. The reproductive characteristics of a species are a result

of the interaction between various endogenous and exogenous factors (Flores and Paula,

2002).

Reproductive biology in brachyuran crabs have been widely investigated mainly

in relation to the monthly occurrence of mature male and females and monthly

distribution of individuals according to size classes and sex. (Colby and Fonseca, 1984;


36

Conde and Diaz 1989(b); Lopez-Greo et al., 2000; Moura and Coelho 2000; Fransozo

and Bertirni, 2002; Gupta, 2012).

Cumberlidge (1999) observed that the gonads of freshwater crabs are located in

the cephalothorax above the hepatopancreas. The male reproductive system consists of a

pair of testes, a pair of vas deferens, two pairs of gonopods (modification of the first and

second pair of pleopods) and a pair of penises. The female reproductive system has been

reported by Cumberlidge (1999); Rostant et al., (2008) to consist of a pair of ovaries, a

pair of ovarian ducts and a pair of spermathecae, which terminate in two genital pores.

After copulation, the ovules develop, resulting in an increase of the ovary size and a

change in color depending on the development stage and species.

According to Hard (1942), female blue crabs in Chesapeake Bay ovulate twice

after maturation and attain sexual maturity in 12-14 months at a size of 5-7 inches. He

has described the macroscopic aspects of the ovaries in this crab during the annual

reproductive cycle and divided the process into various stages. The change in colour of

ovaries is the result of modifications in carotenoid content occurring during the

oogenesis, which play an important role during embryogenesis (Goodwin, 1951).

A smallest adult female blue crab was reported by Fischler (1959) in an ovigerous

condition with a carapace width of 55mm, off the North Carolina coast. Tagatz (1968)

recorded a mature female with a carapace width of 99mm and an immature female with a

carapace width of 177 mm. The shape of the abdomen of maturing females changes from

triangular to rounded and eventually reaches the coxae of the pereiopods. These

variations have been observed in brachyuran crabs such as Libinia emarginata (Hinsch,

1972).

Diwan and Nagabhushnam (1974) in freshwater crab Bartytelphusa cunicularis

observed peak of gonads size in rainy season (June to September) showing the maximum

breeding activity during these months. Histological observations of the monthly gonads

revealed that the ripening of the gonads started during March and extend up to May and

by June the gonads of most of the individuals attained full maturity and spawning begin.

Spawning terminated by the end of September and gonads entered a quiescent period.


37

Changes in colour of ovaries during vitellogenesis is a common charcaterstics in

crustaceans and has been recorded for Macrophthalmus hirtipes (Simons and Jones,

1981) and Crangon crangon Linnaeus, (Haefner and Spaargaren, 1983). According to

Adiyodi and Subramoniana (1983), the colouring of the ovaries is the result of carotenoid

pigments accumulated in the oocytes during vitellegenesis protecting the embryo against

solar radiation. In all ovarian development stages, association between follicular cells and

oocytes were observed. The location of germinative zone being central in brachyurans.

Similar observations have been reported in Portunus sanguinolentus (Ryan, 1967),

Cancer pagurus (Eurenius, 1973) and Callinectes sapidus (Johnson, 1980).

All egg bearing females displayed a rounded abdomen which Christoffersen,

(1988) stated as modification required during incubation of eggs. This widening of

abdomen, he added may serve as an indicator of maturity. According to Shine (1988) in

brachyurans, the females reached morphological sexual maturity at smaller sizes than the

males. This pattern has been stated by Gupta (2012) to indicate the difference between

the sexes in the requirements for reproduction. Both of above workers held that when

females allocate their energy for reproductive purposes, such as spawning and egg

incubation, then they (crabs) tend to mature at smaller sizes than males, who invest their

resources in somatic growth and reach maturity at greater sizes.

The abdominal size and area of pleopod which grew enormously during the

pubertal moult in brachyurans is generally an adaptation to accommodate the maximum

number of eggs produced by a species (Hines, 1991). During the course of gonad

maturation ovaries undergo a sequence of macroscopic changes in its morphology

(mainly in its relative size), which are easily detectable by a naked-eye. (Meusy and

Charniaux-cotton, 1984; Arculeo et al., 1995). The wide variability in carapace shape as

well as abdomen width affects the volume reserved for gonadal development and

consequently spawn size, since the egg mass and the volume of the body cavity usually

present similar allometries (Mantelatto and Fransozo, 1997).

According to Silva (1999), in Panulirus, the presence of highly developed nuclei

indicated that the nuclear components are ready for vitellogenesis. The previtellogenic

oocytes have large peripheral cytoplasmic inclusion body called perinuclear yolk –


38

nucleus complex. The presence of this complex indicates that the oocytes are ready for

vitellogenesis. The macroscopic characterization of the gonads among decapods has been

investigated to determine the onset of the sexual physiological maturity or histological

maturity, taking into account the degree of gonad development. (Costa and Negreiros

Fransozo, 1998; Santos and Negreiros-Fransozo, 1999; Castiglioni and Santos, 2001;

Flores et al., 2002; Castiglioni et al., (2006).

In the freshwater rice field crab, Oziotelphusa sensex senex, Nagaraju et al.,

(2006) induced ovarian growth and vitellogenesis with the introduction of hormones and

suggested that vitellogenesis is a process in which female crabs store nutrients for

subsequent use by developing oocytes. Castiglioni et al., (2007) showed a pronounced

macroscopic differentiation in size and coloration with maturation of the gonads. During

the vitellogenesis, the amount of oocytes in secondary stage increase in the ovary,

resulting in a change in coloration of the gonad. They compared external morphogical

characterstics of the gonads to histological description. They further held that the

macroscopic classification of gonadal stages agree with the modifications of the

reproductive cells (microscopic changes).

In stone crabs, Menippe mercenaria, mature conditions of the ovaries were

determined by gross analysis of the organ and by establishing a color-coded staging

system based upon the color of the ovaries (Gerhart and Bert, 2008). This color-coded

system includes physiologically mature females as having orange gonads, which

indicated that the female was gravid.

Brown (2009) while working on blue crab, Callinectus sapidus in Tampa Bay

described the morphological changes in the female reproductive developmental stages of

reproductive cells. Histological techniques were used by him to determine sexual

maturity of female crabs. Qualitative analysis of gonads were made by describing the

structure of the oocyets and determining the developmental stage of the oocytes from

oogonia to full grown oocytes. Morphological characteristics of ovarian tissue and

oocytes were determined to develop a classification for occyte maturation stages.


39

During maturation process, there are histological and morphological alterations

with change in volume and colour of the gonads. (Rostant et al., 2008). Chen et al.,

(1994) reported colour change in the ovaries of fresh water crabs Eudaniela garmani and

Sinapotamon yangtsekiense , in which transparent ovaries change from white colour to

cream colour, followed by yellow ending up in bright orange, evidenced by the

accumulation of yolk.

Various workers (Castiglioni et al., 2007; Rostant et al., 2008; Santos et al.,.

2009; Souza and Silva, 2009) while conducting the histological analysis of the ovaries

observed a gradual process of oocyte development based on cellular characteristics, viz.,

oogonia, oocytes in initial vitellogenesis, oocytes in advanced vitellogenesis, mature

oocytes, follicular cells and post –ovulatory follicles. All these cells except post ovulatory

follicles types have been found in other species of crabs also. Souza and Silva (2009) in

their work described the morphology of the female reproductive system and the

development of the germ cells of the crab. Goniopsis cruentata. Four stages of ovary

development were defined based on macroscopic and microscope features. The

association of ovarian development stages with biometric measures showed that ovaries

of females with triangular abdomens and a carapace width (CW) of 21.0-29.3mm were in

the previtellogenic stage. Females with rounded abdomens and CW of 26.4-46.1 mm

varied with regard to maturity, the majority being late-stage vitellogenic or mature. The

female reproductive system of G. cruentata was similar to that of most other

brachyurans.

Post–ovulatory follicles are reported for the swimming crabs Callinectes danae

and Callinectes ornatus (Keunecke et al., 2009). Rjeibi et al., (2010) for the first time

reported post ovulatory follicles indicating recent spawning in fresh water crabs. Xue, et

al., (2010) focussed their studies on the developmental changes inside the eggs of the

semi terrestrial fresh water crab. Sinopotamon yangtsekiense from Qiantang river in

China. During development they held that the embryo underwent a series of

morphological changes, followed by decrease in size of yolk mass and color change from

creamy pale yellow to orange and finally grey. The eggs remain attached to the pleopods

in the female‟s abdominal brood pouch during the development.


40

Silva et al., (2012) investigated development stages of male and female gonads in

the freshwater crab Sylviocarcinus pictus through macroscopic and microscopic

(histology) examination. Based on histological description four gonadal stages were

found for females viz., immature, ripening, mature and spawned. Different cells observed

in ovaries were oogonia, oocytes in initial vitellogenesis, oocytes in advanced

vitellogenesic follicular cells and post ovulatory follicles. In males, three development

stages immature, maturing and mature with presence of spermatogonia, spermatocytes

spermatids, spermatozoids and spermatophores were observed. Size at sexual maturity

was 32.3 mm of carapace width for males and 31.5 mm for females. The gonadal stages

observed macroscopically by volume and color were validated through histological

analysis that proved to be useful method for the rapid identification of sexual maturity in

the species.

Herrera et al., (2013) determined the size at sexual maturity in the fresh water

crab Dilcarcinus pagei, from Brazil. The morphological analysis of the gonads was used

to identify and categorize individuals according to their stage of development. The

gonadal maturity was based on the morphology of the gonads which indicated the size at

which the individuals in the population became morphologically mature for reproduction.

Spermatophore formation has been studied for a number of crustaceans using

light microscopy, including the crab species Carcinus maenas by Spalding, (1942) and

Carcinus sapidus by Cronin, (1947). Mota-Alves, (1975) while working on morophology

and histology of the male reproductive system of mangroove crab Ucides cordatusa

observed that testicular lobules are filled with large number of spermatocytes in different

stages of development.

Joshi and Khanna (1982) reported structural and seasonal changes in the testes

and ovary of freshwater crab, Potamon koolooense. The paired H shape testes showed

histomorphological changes during various stages of maturity. A few undifferentiated or

resting spermatogonia supply a new crop of germ cells for the next breeding season.

Spermatogenenetic activity being seasonal, spermatogenesis begins in January- February,

progresses slowly through March, reaching its peak in April-May. Oogonia and young

oocytes develop in the germinal zone, present in the centre of the ovary. The resting or


41

residual oogonia which occur throughout the year divide shortly after ovulation and

supply new crop of germ cells for the next breeding season. Five maturational stages of

ova have been described on the basis of changes that occur in their nuclei and cytoplasm.

They are oogonium, premeiotic oocyte, previtellogenic oocytes, vitellogenic oocyte and

the ripe ovum. Spawning occured during May and June. The weight of the ovaries,

gonad index and ova diameter were minimum in June and reached a maximum value in

April.

Comparisons of the functional morphology of genitalia and subsequent sperm

transfer and storage mechanisms, and the structure of spermatozoa and spermatophores

have been carried out among crustacean taxa very extensively by Bauer, (1986); Tudge,

(1997) and Kronenberger et al., (2004). On the basis of their studies they held that such

studies provide useful information on phylogenetic relationships and evolutionary

divergence, in the Decapoda. Many brachyuran crabs show a bilateral symmetry and H-

shaped male reproductive system consisting of a pair each of testes, vasa differentia ,

penises and gonopods-I and gonopods –II. This arrangement is found in many crabs and

crayfishes (Krol et al.,1992; Cumberlidge, 1999; Lopez-Greco et al., 2007 and Castilho

et al., 2008). Castilho et al., (2008) reported that in Ucides Cordatus and Cardisoma

granhumi, the development of the male gonads occur in two stages viz., immature and

mature, based on the presence or absence of spermatophores in the distal vas deferens

and ejaculatory ducts.

Noro et al., (2007); Erkan et al., (2009) and Santos et al., (2009) observed that in

S.pictus the vas deferens were divided into three distinct regions viz., proximal, medial

and distal. The medial and distal vas deferens are abundantly packed with

spermatophores which are spherical in shape, which is the most common spermatophore

shape among brachyurans. The coloration and cell types in testis followed the pattern

described in the literature for other decapod species. (Castilho et al., 2008; Erkan et al.,

2009 and Santos et al., 2009). They also reported that cell types found in the male

reproductive system were spermatogonia, spermatocytes, spermatids, spermatozoids and

spermatophores, which modified the coloration of the gonads throughout the maturation

process ranging from transparent in immature individuals to white in mature individuals.


42

Sherkhane et al., (2010) described the gross anatomy and histology of the male

reproductive system of Berytelphusa cunicularis. The male reproductive system, they

stated consisted of a pair of testis, a pair of vas deferens ending into ejaculatory ducts and

a pair of penises. The macroscopic anatomy of the testis showed a similar pattern to that

described in other brachyuran species like the presence of paired testis having H shaped

appearance and presence of gonapods. Histologically, the testis consists of large number

of testicular lobes containing spermatogonial cells that differentiate into spermatocytes

and a collecting ducts.

Banumathi, et al., (2013) described the morphology and histological functions of

the testis and vas deferens of Spiralotelphusa hydrodroma. The H shaped testis, they

stated has three distinct regions viz., anterior, intermediate and posterior. Whole anterior

part of the testis exhibits convoluted lobules including seminiferous tubules, the

intermediate part of the testis has a simple columnar epithelium and the posterior part

has a simple squamous epithelium. Gupta (2013) while studying gonadal cycles of

Paratelphusa masoniana from river Chenab reported that these crabs exhibit seasonal

variation on the basis of morphological studies of gonads.

Wojcik and Normant (2014) described for the first time the gonad maturity stage

of Eriocheir sinensis females (carapace width 55.20-78.10mm) collected in the autums

and winters from Southern Baltic Sea. Seventeen females had gonads in the penultimate

stage indicating onset of spawning. Whereas four other females had gonads in the last

stage measuring that they were already carrying eggs. The low salinity permitted mating

and fertilization as well as embryo development in E. sinensis.

Chapter 3

Materials and Methods


43

The studies on ecobiology of Maydelliathelphusa masoniana, a fresh water crab were

carried out at the Department of Zoology, University of Jammu in two phases.

Phase 1: Field studies:

i. Diversity,

ii. Physico-chemical parameters

iii. Population structure.

Phase- 2: Laboratory studies:

i. Morphometric parameters

ii. Nutritional Status

iii. Gonadal development.

3.1 Diversity

Survey was conducted in water bodies of plain and hilly areas of the region for a

period of two years (Jan. 2012-Dec. 2013).


44

Water bodies of plain area: Gho-Manhasan, Gadigarh, Sehi, Sarore

Kheri,Tarnah Mansar and Srunisar.

(Plate -1, Figs 1 & 2)

Water bodies of hilly area: Ban Ganga, Jajjarkotli, Poonch,

Sunderbani, Chenani. (Plate -2, Figs 3&4)

Collection

The crabs were randomly collected from the selected sites by netting or hand

picking by taking the help of local fisherman. Specimens were collected in plastic

containers and brought to the departmental laboratory. Deptt. of Zoology,

University of Jammu.

Morphological analysis was carried out by using normal scale and vernier

callipers. Crabs were identified with the help of identification keys following

Henderson (1893) and Alcock (1909).

Further authentification was done with the help of checklist of Indian fauna of

fresh water crabs by Pati et al., (2013) from Zoological survey of India (ZSI)

Kolkatta.

3.2 Physico-chemical Parameters

Water body of plain area viz Gho-manhasan stream was selected as area of study

during the period of investigation. Gho-manhasan stream which arise from river Chenab

is one of the tributary of Indus river that covers the maximum part of Jammu region of

J&K State. This stream is located at 32.560 N and 74.95

0 E and is used for irrigation as

well as domestic purposes.

Two stations (I and II) were identified for collection purpose, depending upon the

extent of anthropogenic activities.

Station-I

It is the stretch of stream which was found to be moderately disturbed. Villagers

oftenly were seen using water of this station for bathing of both cattle and as well as


45

themselves (Plate-3, Fig 5). Additionally fishermen also used this stretch for fishing

purposes.

Station-II

It is the stretch of stream which was found to have no anthropogenic activities.

This station, located approximately two km away from that of first station, was generally

found to be completely undisturbed as no human physical disturbance could ever be

recorded here (Plate – 3, Fig 6).

Physical Parameters

Temperature: Monthly variations of temperature (air and water) were

recorded from two stations by using mercury bulb

thermometer.

Depth: Water depth of each station was determined with the help

of graduated meter rod.

Chemical Parameters:

Hydrogen ion concentration (pH):

pH of water samples were determined with the help of

portable field pH meter (Hanna).

Dissolved Oxygen:

Dissolved oxygen was analysed from all the stations by

modified Winkler’s method (A.P.H.A., 1985).

Free Carbon dioxide:

It was determined by trimetric method recommended by

A.P.H.A (1985).

Carbonates and Bicarbonates:

Carbonates and Bicarbonates were estimated by methods

employing those suggested by I.S.I (1973) and A.P.H.A.

(1985).


46

Chloride :

Chloride content of water body were analysed by

Argentometric method using potassium chromate as an

indicator (A.P.H.A., 1985).

Calcium:

Calcium was estimated by adopting methodology given by

I.S.I (1973) and A.P.H.A (1985).

Magnesium:

Magnesium was estimated by following methodology as

given I.S.I (1973) and A.P.H.A (1985).

Collection of Crabs

Crabs were collected on monthly basis from Gho-manhasan stream during the

study period (Jan, 2012- Dec 2013). Number of crabs caught in each effort and the total

number of efforts were properly recorded to determine the following.

i. C.P.U.E catch per unit effort has been calculated by following the formula of

Maunder et al., (2006).

Effortsof.NoTotal

CatchTotal.E.U.P.C

3.3 Population Structure

Sex Ratio:

In the laboratory, they were segregated sex wise based on their abdominal

morphology, following which sex ratio was determined by using the formula

propounded by Ali (2004).

CrabsMaleof.No

CrabsFemaleof.NoRatioSex

Size Frequency Distribution:

Based on the carapace measurement, different size classes were constructed

for each sex. (FC1,FC2, FC3, FC4, FC5, and FC6, for females and (MC1, MC2, MC3,

MC4, MC5, MC6 & MC7 for males).


47

3.4 Morphometric Analysis

Morphometric Analysis: Using vernier calliper scale/normal scale crabs were

subjected to morphometric analysis for different morphometric characters as follows.

(Plate 4, Fig A-D)

Carapace Width (CW):

Width between the broader end of carapace denotes

carapace width (Fig: A).

Abdominal Width (ABDW):

Width across midline of the 4th

segment denotes

abdominal width (Fig: B).

Chela Length (CHL):

Length of chela measured from tip of the propodus fixed

finger to the basis of propodus signifies chela length.

(Fig: C).

Chela Depth (CHD):

Measurement across the widest region of the chelipede

palm is the chela depth (Fig: D).

Mean Body Weight (MBDWt):

Recorded on an electronic balance.

In the laboratory, specimens were identified and sexed according to secondary

sexual characters (abdomen morphology and number of pleopods).

Allometric growth/sexual maturity:-

The following variables were measured in the laboratory to the nearest 0.1 cm

using a vernier calliper and total body weight was measured to the nearest 0.1g using a

weighing balance.

a) Carapace width (CW)

b) Chela length (CHL)

c) Chela depth (CHD)

d) Abdominal width (ABDW) in females.

e) Mean body weight of both males and females.(MBDWt.)


48

The morphometric relationship CW/CHL, CW/CHD and CW/MBDWt. for males

and CW/ABDW and CW/MBDWt for females were used to estimate the size of maturity.

The examined variables were subjected to a regression analysis.

In the allometric growth, the carapace width (CW) was used as an independent

variable "y" since it is the most representative dimension of the overall size of the animal

(Hartnoll, 1982) , related to other body dimensions, the dependent variables being "x" in

males , viz chela length (CHL & chela depth (CHD) and Abdomen width (ABDW) in

females including mean body weight (MBDWt) in both the sexes.

Aiming to investigate the occurrence of allometry between the morphometric

variables, specifically those related to secondary sexual characteristics, their values were

logarithrmized (Log y = log a +blog x) and the function Y = a+xb, where x is the intercept

(value of y when x=o), and b the slope of the regression line. The b value indicates the

growth patterns of the analyzed variables, considering three possibilities : b = 1

(isometery) b<1(negative allometry), b>1 (positive allometery) (Hartnoll, 1982). The

statistical significance of b was tested by student t-test.

3.5 Nutritional Status

During the present course of study, only adult male and female crabs, of carapace

width 5-6 cm were selected and the juveniles were again released into their natural

habitat. The analysis was performed for a period of two years (Jan. 2012 to Dec. 2013).

Bio-Chemical Analysis

In the laboratory, mature healthy male and female individuals in the size range of

5 to 6 cm were segregated. Sex wise segregation was done by examining the abdomen

which is triangular incase of male and round in case of female crabs. Monthly collected

samples were subjected to biochemical analysis. After killing, the crabs were dissected

for body meat, all the body meat muscles were weighed on the electronic weighing

machine and their respective weight in grams was recorded.

The organic constituents of each component were determined by standard

methods such as total proteins (Lowry et al., 1951); Lipid (Folch et al., 1956), Moisture


49

and Ash (Standard method of AOAC, 1999). The results were expressed on dry weight

basis.

The organic constituent of each component were determined by standard methods such

as:

1. Estimation of Protein Content: Protein content of the body parts of

M.masoniana was estimated following Lowry et al., (1951).

2. Lipid estimation: The total lipid content was determined by the method of Folch

et al., (1956)

3. Estimation of Moisture content: Moisture content was determined by standard

method of AOAC, 1999.

4. Estimation of total Ash Content: Ash content was estimated by heating the

samples in a muffle furnace at 5500c for about 3 hours. (AOAC, 1999).

Crude Protein:

Preparation of tissue homogenate:

To a 10mg of sample, 1 ml of 1N NaOH was added for protein extraction in water

bath of 30 minutes. Thereafter, it was cooled at room temperature and neutralized with

1ml of 1N HCL. The extracted sample was centrifuged at 2000 rpm for 10 minutes, and

an aliquot of the sample (1ml) was further diluted with distilled water (1/9v/v) and was

preserved for further analysis.

Reagents required.

a) BSA (Bovine Serum Albumin) stock solution (1mg/ml)

b) Analytical reagents.

i. Sol. A: 50ml of 2% Sodium carbonate mixed with 50ml of 0.1 NaOH

solution (0.4g in 100ml of distilled water).

ii. Sol. B: 10ml of 1.56% CuSO4 solution mixed with 10ml 2.37% Sodium-

potassium-tartarate solution.

iii. Prepare analytical reagent by mixing 2ml of sol. B with 100ml of Sol. A.


50

iv. Folin-ciocalteau solution (1N):

Dilute commercial reagent (2N) with an equal volume of water on the day of use

(2ml of commercial reagent + 2ml distilled H20).

Principle:

The phenolic group of tyrosine and tryptophan residues (amino acid) in a protein

will produce a blue purple coloured complex with maximum absorption in the region of

660 nm wavelength with Folin-ciocalteau reagent which consists of sodium-tungstate

molybdate and phosphate. Thus, the intensity of colour depends on the amount of these

aromatic aminoacids present and will thus vary for different proteins. Most proteins

estimation technique use BSA universally as a standard protein because of its low cost,

high purity and ready availability. The method is sensitive down to about 10μg/ml and is

probably the most widely used protein assay despite its being only a relative method,

subject to interference from Tris buffer, EDTA, non-ionic and cationic detergents,

carbohydrate, lipids and some salts. The incubation time is very critical for a reproducible

assay. The reaction is also dependent on the pH and a working range of pH 9 to 10.5 is

essential.

Procedure:

Different dilutions of BSA are prepared by mixing stock BSA (1mg/ml) and

distilled water in the test tubes. The final volume in each of the test tube is 5ml.

The BSA range is 0.05-1 mg/ml.

From these different solutions, pipette out 0.2ml protein solution to different test

tubes and add 2.5ml of alkaline CuSO4 (analytical regent). Mix the solutions well.

This solution is incubated at room temperature for 10 minutes.

Then add 0.5ml of reagent Folin-ciocalteau solution to each test tube and incubate

for 30 minutes. Set the calorimeter at zero with blank and take the optical density

(measure the absorbance) at 660nm.

Plot the absorbance againt protein concentration to get a standard calibration

curve.


51

Check the absorbance of unknown sample and determine the concentration of the

unknown sample using the standard curve plotted above.

For the diluted sample, 1 ml was taken and treated with 2.5 ml of mixed reagent

(carbonate-tartarate-copper) and 0.5ml of 1N Folin’s regent. After 30 minutes, sample

absorbency was read at 750 nm using spectrophotometer. The results were expressed as

percentage.

Lipids:

Lipid occur in tissues in a variety of physical forms, but the complex lipids are

usually constituents of membrane, where they occur in close association with such

compounds as protein and polysaccharides, with which they interact by hydrophobic and

vender vaals. Various solvent or solvent combinations are used as extractants but mixture

of chloroform-methanol (2:1 by volume) has been found to be ideal solvent as per Folch

et al., 1957 method.

Chemicals required:

1. Methanol

2. Chloroform

3. Sodium Chloride

Procedure:

Sample was weighed (1-2g) accurately and mascerated using mortar and pestle

adding 5ml of methanol-chloroform mixture (2:1). After proper masceration put mixture

in a measuring cylinder and added to it 20 ml of chloroform-methanol mixture again.

Mixture was kept overnight. It was then filtered using Whatman filter paper and the

filtrate was collected. The residue was collected and the procedure was repeated by

adding 25ml of chloroform methanol mixture and kept overnight. Supernatant was

collected. To pooled supernatant added 1/5th

volume of 0.9% NaCl solution to the

collected filtrate. Put the supernatant in a separating funnel. Mixture partitioned into two

layers of which the lower was composed of chloroform-methanol containing virtually all

of the lipids while the upper phase contained same solvents with non-lipid contaminants.

Lower phase was collected in a pre-weighed beaker. It was then dried on water bath at a


52

temperature of 400C. Took the final weight and percentage lipid was calculated by using

the formula.

Calculation:

W

100WWLipidofage% 12

Where,

W = Wt of the sample

W2 = Wt. of beaker + chloroform methanol extract

W1 = Wt of empty beaker

W2W1 = Wt of crude lipid.

Moisture:

Sample when dried in an oven at 1050 ±1

0c to a constant weight, the loss of

weight of a sample is an index of moisture content. For estimation of moisture, sample

weighing 0.5 to 2.0 g was taken in triplicate, kept into pre-weighed aluminium dishes.

Dried sample (with their lids removed) were then placed in an oven and dried for 11 hrs

at 1050 ±1

0c. After this, lids were placed over dishes while still in oven and then

transferred to dessicator. After a sufficient period of time, when room temperature is

attained, again weight is taken. Loss in weight gives the moisture content.

Moisture content of the sample (%) = WW

100WW

1

21

Where,

W = Wt. of empty dish.

W1 = Wt. of dish plus sample before drying.

W2 = Wt. of dish after drying.

Ash Content:

Ash is the organic residue remaining after sample is ignited to carbon free, usually

at a temperature not exceeding red hot. Sample used for the determination of the ash

content, was weighed accurately (1-2 gm) in triplicate in silica crucibles. Crucibles were

than kept in a muffle furnace raising the temperature slowly to 6000C. Sample is ignited


53

for four hours at 6000C. After heating, crucibles are transferred directly to dessicator,

cooled and weighed. After drying, the left over residue represents the ash content which

was calculated by using the formula.

%ash = 100WW

WW

01

02

Where,

W1 = Wt of crucible plus sample

W0 = Wt. of empty crucible

W2 = Wt of crucible plus sample after ignition

W1-W0 = Wt of sample

W2-W0 = Wt. of residue representing ash content

3.6: Gonadal Development

In the laboratory, crabs were anaesthetized by deep freezing them for 30 minutes.

They were then washed and segregated sex wise. The size of male and female crab i.e.

carapace width (CW) was measured with vernier calliper scale/normal scale. The study is

based on a total of 48 specimens per year (24 each sex). After making macroscopic

observations they were dissected out for microscopic studies in order to observe seasonal

variations in gonads on monthly basis.

The gonadal developmental stages for both sexes of M. masoniana were

investigated by following methods.

(a) Macroscopic examination of the consistence, volume and coloration of the ovaries

and testis in relation to thoracic cavity (Mantelato & Fransozo, 1999)

(b) Microscopic examination of cell types (Castilho et al., 2008)

For macroscopic and microscopic analysis 24 gonads of each sex were randomly

selected from sexually mature specimens ranging in CW from 3.5 to 5.5 cm.

For macroscopic analysis, the shape, size and coloration of gonads were analyzed

and photographed and classified as immature, maturing, mature and spawned following

the pattern of Haefner (1976) and Fransozo et al., (2002).


54

For histological description, gonads were fixed in Bouins fixative for 24 hrs, at

200C. After post fixation treatment gonads were kept in 70% alcohol and subsequently

dehydrated through alcohol series (70% to 100%), diaphanized in xylol, infiltrated and

embedded in paraffin. Finally, the sections were prepared using a microtome (3 to 4 μm)

and stained using haematoxylin-eosin. The thin sections of ovaries and testis were

analyzed with an optical microscope and photo micrographed using Olympus CH20

iBIMF attached with Sony SSC-DC378 P Camera.

Photography:

Photography of various survey sites and specimens were conducted by Sony

Cyber-short DSC-W310.

Statistical Analysis

Graphs & Correlations : MS Excel

Regression/t test : SPSS

(Statistical Package for Social Sciences)

version 18.0.

Chi-Square Test (Chi.sq) : Statistical Version 7 software is used for

data analysis.

Chapter 4

Results and Discussion


55

4.1 Diversity of crabs, their identification and taxonomic revision.

Two species of freshwater crabs belonging to different genera have been recorded

during present study period. (Table 4.1.1: Fig 1 and 2) (Plate 5)

Species: I

Taxonomic position of Maydelliathelphusa masoniana (Henderson, 1893)

Phylum-Arthropoda

Subphylum-Crustacea

Class-Malacostraca

Order-Decapoda

Sub order-Pleocyemata

Infra order-Brachyura

Super family-Gecaricinucoidea

Family-Gecarcinucidae

Genus-Maydelliathelphusa

Species –masoniana


56

I. Maydelliathelphusa masoniana (Henderson, 1893) Fig 1 : (i) a, b & c. (Plate 6)

Family: Gecarcinucidae Rathbun, 1904.

Specimens examined: 24 males and 24 females per year.

Colour: All the specimens examined are characteristic in colour with dark chocolate

brown, shining carapace on dorsal side. The ventral surface of carapace being light brown

in colour. (Fig 1: (i) a & b).

Size: Species attains the size upto 7cm of carapace width (C.W), especially in males.

Whereas females of the same species are smaller with size upto 6 cm. of carapace width

(cw). (Fig 1: (i) a & b). (Table 4.1.2)

Heterochelous chelas: Heterochelous Chelas have been observed to be present in both

the sexes with heavy, unequal and bigger size chelipeds in males than females. Cutting

edges of chelas are armed with teeth of variable size.All the teeth and tips of both fingers

are dark brown in colour . (Fig 1: (i) c).

Habits: As observed by their random gut content analysis , species is omnivorous,

feeding on algae, water plants, detritus, and insect larvae. Maximum crab population is

recorded during months of June-July and December-January, thus exhibiting them as

biannual breeders. Mature adults, both male and females are witnessed in collection

during advancing summers (April-June) and secondly during approaching winters

(October-December) whereas juveniles were reported in rest of months.

Habitat: Abundant in shallow water bodies such as paddy fields, canals, ponds etc. and

prefers to construct burrows closer to water level in clay soil to escape from dry seasons.

Distribution: Water bodies of plain areas of Jammu region (J&K) viz. Gho-manhasan,

Gadigarh, Sehi, Sarore.

Species: II

Taxonomic position of Himalayapotamon emphysetum (Alcock 1909)

Phylum - Arthropoda

Subphylum - Crustacca

Class - Malacostracca

Order - Decapoda


57

Sub order - Pleocyemata

Infra order - Brachyura

Super family - Potamoidea

Family - Potamidae

Genus - Himalayapotamon

Species - emphysetum

II. Himalayapotamon emphysetum (Alcock 1909) Fig: 2 (ii) a, b & c. (Plate: 7)

Family: Potamidae

Specimens examined: 12 males and 12 females per years

Colour: All the specimens examined have reddish orange to greyish brown coloured

dorsal surface, which is rough but not shining. The ventral surface of carapace being pale

white in colour. (Fig 2: (ii) a & b).

Size: Specimens of this species are of moderate size, males reaching a size upto 5 cm of

carapace width (cw). Females being smaller than males with size range of 4.5 cm of

carapace width (cw). (Fig 2: (ii) a & b). (Table 4.1.3)

Heterochelous Chelas: Heterochelous chelas used for grasping defence and sexual

signaling are medium sized, less serrated but have a peculiar feature of purple or blue

with orange coloration at their tips. (Fig 2: (ii) c).

Habits: Omnivorous, feeding on both plant and animal matter. Crab population of the

species records presence of all age groups viz. adult male/females and juveniles all the

year round thus exhibiting them as continuous breeders. The species is seen in large

numbers wandering around on land particularly soon after rain for food which is

abundant at this time on land then in water.

Habitat : Abundant in running water condition under stones and boulders present in

water bodies of hilly areas. The species is confined either to high gradient streams or to

more or less sluggish rivers.

Distribution: Banganga, Jajjarkotli, Poonch, Sunderbani, Chenani.


58

Discussion

During the present investigation, two species of freshwater crabs have been recorded viz

Maydelliathelphusa masoniana Henderson (1893) and Himalayapotamon emphysetum

Alcock (1909) from water bodies of Jammu region. (Table 4.1.1).

The former species Maydelliathelphusa masoniana Henderson, (1893) is a

taxonomic revision of Paratelphusa masoniana Henderson (1893) reported from

Gadigarh stream, Dutta (1978), Gho-manhasan stream, Gupta (2012) of Jammu region

(J&K). The taxonomic position of later species Himalayapotamon emphysetum Alcock,

(1909) has been reported from the water bodies of hilly areas viz. Banganga, Jajjarkotli,

Sunderbani, Poonch and Chenani of Jammu region.

The Global position showing the diversity of fresh water crabs indicates that fresh

water crab taxonomy is regarded as still being in its discovery phase. Yeo (2007). There

are currently total of 238 genera including 1,306 true fresh water crabs species

accommodated in six families. (Fig: 3) The oriental region is taxonomically most diverse

part of the world with over 900 species of fresh water crabs in 154 genera and 2 families

viz Gecarcinucidac and Potamidae Yeo et al., (2008), Cumberlidge et al., (2009).

According to Dai (1999), the family Potamidae reaches its greatest diversity in

oriental region hosting about 450 species. On similar lines Ng et al., (2008) reported that

family Gecarcinucidac has a total of 345 species and being diverse in oriental region viz.

North-East India, Myanmar, Sri Lanka. Cox, (2001) as also depicted in (Fig: 4; Map -I)

The taxonomic position and confinement of Maydelliathelphusa masoniana under

family Gecarcinucidac in water bodies of plain areas and Himalayapotamon emphysetum

under family Potamida from water bodies of hilly area gets authenticated by work of

Cumberlidge (1999). According to him family Potamidae is derived from Greek word

‘Potamon’ meaning river where as family Gecarcinucidac refers to ‘land crabs’. In some

parts of tropics, some species have moved out of rivers, streams and lakes and have

colonized nearby land.

The origin of gecarcinucoid crabs in Africa and their reaching South Asia via

transoceanic dispersal was suggested by Klaus et al., (2006). Whereas, the dominance of


59

gecarcinucoid crabs in Indian peninsula along with presence of large number of potamids

in continental Asia was reported by Ng and Rodriguez (1995).

Further Brandis and Sharma (2005) while dealing with Maydelliathelphusa

examined many specimens from Nepal and northern India. They concluded that this

species is distributed along with Ganges and Brahmaputra valleys ranging from Punjab to

Nagaland. It predominantly inhabits the lowland banks of rivers, reaching higher altitudes

upto 3500m in Himalaya. On similar lines Ng et al., (2008) elevated Maydelliathelphusa

to the genus level and listed five species viz Maydelliathelphusa edentate, lugubris,

falcidigitis, harpax and masoniana.

The presence of gecarcinucoid crabs in Central and Northern India can be

authenticated from the work of Alcock (1909, 1910) who reported 12 species of

gecarcinucid crabs from these areas. From the Himalayas, the first mention of fresh water

crabs was by Wood-Mason (1871) for Potamon edwardsi and P.andersoniaum from the

eastern Himalayas. Rathbun (1904-1906) placed these species in the genus Potamon.

Whereas six years later Alcock (1909-1910) re-described the species of Wood-Mason as

Potamon atkinsonianum and established several sub species as Potamon emphysetum

(Alock, 1909). For taxonomic classification, the characters like morophology of male

copulatory organs were taken into consideration.

Based on these characters, Pretzmann (1966b) established the subgenus. Potamon

(Himalayapotamon) for P. atkinsonianum and added some new sub species. Thus

Himalayapotamon has been given the generic status to include Himalayapotamon

atkinsonianum and Himalayapotamon emphysetum. Later Bott (1970) too, authenticated

the logic of including Himalayan species of fresh water crabs to the genus Potamon. The

distribution of Himalayapotamon in western and central Himalayas and being endemic to

this region was exhibited by Brandis et al., (2000) who held that the distribution pattern

of two species of Himalayapotmon exhibits one species viz. Himalayapotamon

atkinsonianum occurring in east part and other species viz. Himalayapotamon

emphysetum occurring in west part (Fig: 5; Map: 2). Later the genus Himalayapotamon

emphysetum has been reported from North Western India, Punjab, Himachal Pardesh,

Western to Central Nepal as presented in (Fig:5; Map: 2).


60

Present report is therefore, updates our existing knowledge authenticating the

presence of Himalayapotamon emphysetum in the hilly stretches of Jammu region viz

Sunderbani, Poonch, Jajjar and Ban Ganga streams and Maydelliathelphusa masoniana

in the plain stretches of Gho-manhasn, Garigarh, Sehi and Sarore. (Table: 4.1.1; Fig 6:

Map 3)

On the similar lines Maydelliathelphusa masoniana has been reported from the

fresh water bodies of plain areas of Jammu region (J&K), North India. (Table 4.1.1.)

Whereas, Himalayapotamon emphysetum has been reported from the hilly areas of the

Jammu region. (Fig 6; Map 3)


61

4.2 Physico chemical parameters in relation to abundance of Maydelliathelphusa

masoniana

Data on C.P.U.E on monthly basis for two different stations is depicted in table

4.2.1. A look at the table revealed that values of C.P.U.E never remained constant but

fluctuated seasonally as well as at two different stations. It was clearly evident that value

of C.P.U.E is comparatively higher at station II as compared to station I. Maunder et al

(2006) while studying the relationship between C.P.U.E and abundance of individual

stocks held that C.P.U.E varies with (a) efficiency of a fleet (b) species being targeted

by a fleet and (c) environmental factors. Since there was only one species of crab in

stream (target species), therefore, both factors (a) and (b) were constant but

environmental factors i.e. physico-chemical parameters and anthropogenic disturbances

(bathing by human and cattle, in and around the stream) varied to great extent at two

stations seasonally.

Discussion

The crustaceans are highly sensitive to pollution and their distributions are

strongly influenced by physico-chemical parameters. The variations in physico-chemical

parameters influence crustacean’s abundance and life cycle. Species exhibit year to year

fluctuations in abundance as physical, chemical and biological features can strongly

influence abundance.

Temperature

Temperature is a limiting factor in aquatic environment and considerably affects

various metabolic activities, growth, oxygen consumption, reproduction molting, survival

distribution and migratory behaviour of crustaceans. The air and water temperature were

found to go more or less hand in hand.

During the study period (Jan 2012 –Dec 2013), the air temperature at two stations

fluctuated between 160C to 40

0C with maxima in June viz. 40

0C and 38.5

0C and minima

in the month of December viz. 160C and 12

0C. Like air temperature, water temperature

showed similar trend at two stations with fluctuation between 9.50C to 34

0C with

maxima in June viz. 340C and 32

0C and minima in the month of December viz. 9.5

0C

and12.50C during two years of study period. (Tables and Graphs 4.2.2 and 4.2.3). In the


62

present investigation, total crab population showed a direct positive correlation with

water temperature at both the stations viz. values being r=.003147 and r=0.009885 for

first year and r=0.24342 and r=0.2113549 during second year of study period (Table-

4.2.4) That the existing temperature range fall within the tolerable limits of crab is

indicated by their respective values of CPUE (Table 4.2.1) and gets authenticated by

those made by Diwan and Nagabhushanam (1976) who on the basis of their

investigations on the heat tolerance in crab Barytelphusa cunicularis, concluded 34.5 ±

0.50C to be the critical limiting temperature for the said species.

Apart from survival, temperature also play an important role in breeding. It was

observed that juveniles started appearing in the collection during March-June there by

indicating a rise in temperature to be a stimulant for hatching. That the hatching bears a

positive co-relation with increasing temperature further gets authenticated by the

observation already on record by Anger (1991) who while studying the effect of

temperature on larval development of Chinese mitten Crab, Enocheir sinesis, reported

that development from hatching to metamorphosis occurred at temperature more than

120C. In this context, the observations made by workers like Manohar and Qureshi

(1996) who studied the prawn ecology also witness a direct correlation with prevailing

water temp (r=0.1111). Though very little variations in temperature were evident at the

two study stations, yet the variations in population structure as indicated by C.P.U.E were

remarkable. This can certainly be attributed to more anthropogenic activities at Station I

as compared to Station II (Tables and Graphs 4.2.2 and 4.2.3).

pH

In decapods, pH influences the metabolism, physiology and maturation process.

Presently pH value range between 6.7 to 7.4 at both the stations with maxima of 7.4 at

station II during second year of study period. This high value of pH in the range of 7.4 at

station II simply implies that water at station II is slightly alkaline and more basic as

compared to station I (Tables and Graphs 4.2.2 and 4.2.3). Quite interestingly, crabs also

followed the same trend of their population abundance viz. station II > station I as

indicated by C.P.U.E . Thus the relationship of pH to the crab abundance registered

significant positive correlation at both the stations with the values being r=0.449495 and

r=0.338326 for first year and this trend got authenticated by the observation values


63

recorded during the second year of studied period r=0.329087 and r= 0.25577 (Table

4.2.4).

Present observations get support from the work of Das and Sahoo (1997) who

held alkaline pH to be associated with more number of crab species upto a particular

extent after which further increase in pH leads to a decrease in the number of species.

Less crab population at station I can be attributed to slight acidic pH, which appears to be

the main cause for decline of crabs at station I than at station II. This is in accordance to

EPA (1980) that accepted water quality criteria indicating a pH of less than 6.5 units to

be harmful for many aquatic species. Moreover, in this context a strong support can be

drawn from the findings of Sarah (2007) that pH have great effect on immunity of crabs,

higher the pH, more immune crabs are and a fall in pH make them vulnerable to

infection. At lower pH 6.7 the activity of phenol oxidase enzymes which helps to provide

protection against infection in arthropod get suppressed. This very clearly indicates that

pH , like other parameters have marked influence on distribution /abundance of crabs.

Depth

During period of investigation the depth ranged between 14 to 42 cm at two

stations, with station I observed to be more deep with maximum depth of 42 cm as

compared to station II with maximum depth of 32 cm (Tables and Graphs 4.2.2 and

4.2.3). Crab abundance at station I with more depth showed a negative correlation, the

values being r=-0.024061 and r=-0.00435. Station II which was observed to be less deep

showed positive correlation with values r=0.015695 and r= 0.212359 during two years of

study period (Table –4.2.4). A similar relationship of depth with crab population has been

highlighted by Marijnisson (2008) who stated that crab density decrease with increase in

depth of water body.

Dissolved Oxygen (DO)

Oxygen is the prime requirement of any organism, dissolved oxygen being one of

the important parameter in water quality assessment and is an index of the biological and

physical processes operating in water.

During present course of study, the values of dissolved oxygen at two stations

were observed to show great variations ranging from 3.2 to 9.4 mg/ litre. At station I DO


64

fluctuated between 3.2 to 7.0 mg per litre and at station II, the values ranged between 5.2

to 9.4 mg/litre. It was also observed that at station I, DO level remained below 5.5

whereas at station II, DO levels remained above 5.5 for most of the period of

investigation, thus indicating station I to be more degraded than station II (Tables and

Graphs 4.2.2 and 4.2.3). A very strong positive correlation of DO and crab abundance

could be recorded at stations (I and II) with values (r=0.095074 and r=0.174293) during

first year and values (r=0.025618 and r=0.119852) during II year of study period (Table

4.2.4). Similar correlation were established by Manohar and Qureshi (1994) in prawn

with ‘r’ value being r=0.0778 with respect to dissolved oxygen.

Cheng (2003) while studying Haliotis diverscolor reported parameters like

haemolymph, osmolalrity and sodium balance at very low concentration at low DO of

3.08 mg/l. Values lower than this might result in acid base imbalance leading ultimately

to acidosis for short term period which further may limit distribution of crabs, if low

concentration of DO continues to prevail.

Present observations indicated that station I with DO less than 6mg/l appeared

degraded because of anthopogenic activities as compared to station II. This is supported

by findings of Egemen and Sunlu (1999) according to which the minimum DO should not

be <5.0mg/l for survival of aquatic life in fresh water ecosystem. This is further

supported by CPCB (2005) that water bodies with DO at 6mg/l or high are categorized as

class I and less than 6mg/l is categorized as low class or polluted.

FCO2

FCO2 usually maintains an inverse relationship with pH but a direct one with that

of temperature. Tables 4.2.2 and 4.2.3 very clearly, reflect that FCO2 fluctuates between

4.0-12mg/l at both the stations. At station I the FCO2 fluctuated between 4.0 to 12.0 mg

per litre and at station II FCO2 ranged between 4.2 to 7.6mg/liter indicating high levels

of FCO2 at station I as compared to station II. With respect to crab abundance, FCO2

showed negative correlation at station I with ‘r’ values being -0.396196 and -0.01978

during Ist and IInd year respectively. A positive correlation however, at station II with

‘r’ values of 0.13258 and 0.394468 during 1st and IInd year of the study period (Table

4.2.4) has been witnessed.


65

At station 1, FCO2 value was high during summers at high temperature (May-

June) hovering between 10.0-12.0 mg/l but was inversely related to pH value ranging

between 6.8-6.9. Since FC02 has great bearing on pH, therefore it can have strong

influence on the crab population. Increased CO2 related increase in pH at station may be

responsible for resultant decline in crab population by disturbing acid base balance of

ecosystem. This can be authenticated by findings of Spicer et al., (2007) who also stated

that CO2 related acidification leads to disturbance of acid base balance in velvet

swimming crab Neuora puber.

Further at station I, anthropogenic activities add to the rate of decomposition and

as a consequence CO2 levels also mark an increase which further lead to a decline in

crab population at this station. At station II, a positive correlation of FCO2 with crab

abundance can be attributed to low values of FCO2 during summer (6.2-7.4 mg/l) and

more basic values of pH (7.1 and 7.2) as compared to station I showing negative

correlation with crab abundance because of high values of FCO2 and less basic values of

pH. (Tables and Graphs 4.2.2 and 4.2.3). Further no anthropogenic activities were

witnessed at station II. This comparison strongly indicated role of FCO2 in crab

distribution which was much pronounced at Station II than at Station I.

Chloride

Chlorides occur naturally in all types of water. High concentration of chloride is

considered to be the indicator of pollution due to organic wastes of animal or industrial

origin. Chlorides are troublesome in irrigation water and also harmful to aquatic life.

The levels of Cl- ranged from 23.6 to 56.8mg/l and 20.0 to 40.9 mg/l at station I

and station II respectively (Tables and Graphs 4.2.2 and 4.2.3). It was observed that

station II had low chloride levels than station I indicating that increase in salinity at

station I resulted in a decrease in crab population as compared to station II where the

levels of chloride are less than station I. Since chlorides help maintaining the fluid and

acid base balance, therefore, the increased salinity of water cause acidosis and serve as

limiting factors for the distribution of crabs. Whitely et al., (2001) too held that an

increase in salinity disturbs acid base balance and rather cause acidosis in crabs and thus

support the view that concentration of Cl- does influence the distribution of Crabs. Thus


66

Crab population of present investigation exhibited a negative correlation with chloride at

station I with values (r=-0.02784 and r=-0.08355) and with the values (r=-0.00447 and

r=-0.058066) for station II during first and second year of study period respectively

(Table 4.2.4). Manohar and Qureshi (1996) also reported similar relationship of prawn

population and chloride (r = -0.0598).

It was also noticed during present study that high values of chloride content at

station I were due to anthropogenic activities which in turn led to a decline in crab

population as also been earlier highlighted by Khare et al., (2007) according to which Cl-

as an important indicator of pollution. On the other hand absence of anthropogenic

activities along with low level of Cl-

at station II indicated more congenial habitat for

fresh water crabs.

Ca2+

and Mg+

Calcium and Magnesium are very vital components so far as crabs are concerned

because they have their special requirement for these ions during the formation of

exoskeleton (Neufeldt et al., 1994).

Though in comparison to Ca++

, Mg++

is required in very small concentrations.

Mg++

fluctuated between 40.2 -64.7 mg/l at station –I and between 34.0 -56.9 mg/l at

station –II showing a positive correlation with crab abundance at two stations with values

(r=0.495669 and r= 0.602844) during first year and with values (r=0.0150749 and r=

0.496578) during second year of study period (Tables 4.2.2, 4.2.3 and 4.2.4).

Whereas the levels of Ca2+

has been observed to be high at station II viz 20.0 to

54.5 mg/l as compared to station I (22.8 - 42.2 mg/l). Higher levels of Ca2+

at station II

exhibit a positive correlation (r = 0.056325) with respect to crab abundance at this station

but station I indicated negative correlation ( r = -0.326441) (Table–4.2.4). Present

observations get support by the findings of Neufeldt et al., (1994) who while studying the

effect of calcium on its post moult uptake in blue crab Callinectes sapidus reported that

rate of calcium uptake in these crabs have direct relation with availability of Ca2+

in water

body as their calcium requirement in considerably high during ecdysis. At station I,

negative correlation appears to be the indicator of the fact that extremely low calcium

possibly must have led to incomplete calcification of developing crabs. These crabs,


67

because of low calcium availability have poor exoskeleton and became highly vulnerable

to predation and cannibalism which ultimately have resulted in limiting their distribution

at station –I. In this context, findings of Rayhanen, (1962) that in crustaceans, incomplete

calcification lead to prolonged period of soft exoskeleton, making them more vulnerable

to predation, very strongly justifies the less crab abundance at station –I

In comparison to Station I, Station II with high value of Ca2+

and positive

correlation with more abundance of crab population comprising of both the juveniles and

adults strongly indicated that calcium may limit the distribution and success of

crustaceans in soft water localities.

Carbonates (CO32 ) and Bicarbonates (HCO

-3)

In water, CO2 exist in three different forms viz FCO2, bound CO32-

and half

bound form (HCO3-

). During present studies bicarbonates have been observed to be

present at two stations but (CO32) could not be ever recorded at any station as FCO2 was

present. Role of bicarbonates as buffer in a water body is of utmost significance

particularly when it comes to support biotic communities. It appears that bicarbonates by

regulating pH in turn control the crab population abundance and same may be true for

presently studied stream as well.

At station I value of HCO3 ranged between 430.3-886.3 mg/l with a positive

correlation (r=0.183321 and r= 0.124026) during first and second year of study period

respectively. At station II, HCO3 ranged between 400.2-752.0 mg/l showing a negative

correlation with values r=-0.24529 and r=-0.16536 during first and second year of study

period respectively (Table 4.2.2, 4.2.3 and 4.2.4). Our observations indicated that water at

station II is more productive than water at station I. It was observed that at both the

stations the level of HCO3 was on higher side during summer and monsoon period.

Present observations are in line with those made by Zuber (2007) regarding increase in

value of bicarbonates in summer seasons. Verma (2009) too, emphasized maximum value

of bicarbonates during monsoon because of entry of rain water that is rich in CaCo3

which in turn increase the bicarbonate concentration in study area.


68

4.3: Population structure of M.masoniana from natural habitat

4.3.1: Sex Ratio

A total of 592 crabs were caught during the study period, 310 (52.36%) turned to

the females against a total of 282 (47.63%) males Table 4.3.1, Fig: 7. Sex ratio therefore,

comes to be 1:1.09. Sex ratio, however never remain consistent, rather it exhibited

seasonal variations. As evident from Table 4.3.1 and Graph 4.3.1 females were more

abundant then males in the month of June and July and males were dominant in month of

Dec. However, Chisquare test (Chi.sq =7.8259; df=11; p-value < 0.7288) showed that sex

ratio may be considered independent of the catching month.

4.3.2: Size Frequency Distribution

On the basis of carapace width, different size classes were constructed for each

sex (Table 4.3.2 and 4.3.3). Size of male crabs ranged from 2-3cm CW to 6-7 cm CW. In

females, the size range was observed to be 2-3cm to 5-6 cm CW. No female was recorded

in the class size of 6-7 cm CW. (Graph 4.3.2 and 4.3.3)

From the study it is evident that comparatively males maintain large size than

females. Crab population have been observed to comprise primarily of adult individuals

(size>3cm CW). Juveniles/ small crabs were rarely spotted and that too in the ending

August and March, months.

4.3.3: Breeding season

From the present study as depicted in the table 4.3.2 and 4.3.3 the crab

M.masoniana exhibit two peaks in breeding season viz. June-July and December-

January. It is during these months that sexually mature males and females with mean of

5-6 cm CW and 4-5 cm CW respectively were found maximum in the collection with

highest of 17 males and 16 females, in the month of December.

Discussion

4.3.1: Sex Ratio

The overall sex ratio differed from expected mean value of 1:1 during the months

of June-July and December-January which happen to be the breeding season of this crab.

Several causes may lead to this discrepancy such as differences between sexes in growth


69

rate, differential migration and mortality. According to Werner (1972) sex ratios differing

from 1:1 are widespread among crustaceans. In M.masoniana, females tend to be more

abundant than males (Table 4.3.2 and 4.3.3.). The present study is in accordance to those

recorded in hermit crab, where females tend to be more abundant than males (Ameyow-

Akumfi, 1975; Martinelli et al., 2002).

Our observation is also in accordance with the findings of Czernijewski and

Wawrzynailew (2006). In Chinese mitten crab Eriocheir sinesiss where the overall sex

ratio of 1:0.80 differed from the expected 1:1 ratio, with males dominant in month of

December and female being abundant in the months of October and November.

Fluctuation in the sex ratio can be attributed to various factors (i) migration of male crabs

towards sexually mature females during breeding season. (ii) burrowing habit of females

during breeding season for incubation of eggs and protection of brood. Similar findings

have also been reported in fiddler crabs and Uca crenulata (de Rivera, 2003) wherein he

reported the burrowing habit of ovigerous females to be one of the major factor

contributing towards the disturbed sex ratios.

Present findings also get support from the work of Ali et.al, (2004) in Scylla

Serrata of Sunderbans mangrove where overall sex ratio of male to female was found to

be 1:0.94 and fluctuations on monthly basis could also be observed. Sallam (2005) too

reported fluctuation in sex ratio of Dotilla Sulccats from 1:1 with more males than

females. Our study get strengthened from report of Lawal-Are (2010) who studied sex

ratio in blue Crab, Callinectes amnicola to be 1:0.96 exhibiting fluctuating pattern and

females being abundant than males in rainy season.

4.3.2: Size Frequency Distribution

M.masoniana is sexually dimorphic with males being larger than females viz

females with maximum of 5-6 cm CW and males reaching upto 6-7 cm of CW (Table

4.3.2 and 4.3.3). The present study exhibited that the females reached morphological

sexual maturity at smaller size 4-5 cm CW than males of size 5-6 cm CW. This

observation is consistent with the pattern proposed by Shine (1988) for brachyurans who

held that females allocate their energy for reproductive purpose such as spawning and

egg incubation and therefore, tend to mature at smaller sizes than males, who invest their


70

resources in somatic growth and reach maturity at greater sizes. Present observations are

also in accordance with the findings of Jivoff (1997) who held that in male blue crab,

Callinecles sapidus, males attain maturity at size bigger than females.

On similar lines, Diaz and Conde (1989) while working on size frequency

distribution exhibited that males reaching larger size than females, characterizes a

dynamic equilibrium for a certain population with slight monthly variations and

differential mortality rates. From the present study, it is evident that three factors can help

to determine the size of local population of M. masoniana (i) the difference in energy

available for growth, with males growing more because they don’t spend energy in egg

production; but use their energy for structural metabolism (ii) the large reproductive

effort exhibited by males may be due to their ability to fertilize more than one female.

(iii) Males of large size have greater chance of obtaining females for copulation as a

function of intra-specific fights (Abrams, 1988).

From the Tables 4.3.2 and 4.3.3 it is evident that the no of females of class size 4-

5 cm CW were numerous than males of same class size. However, males were reported to

be dominant in larger class size of 5-6 cm CW (Table 4.3.3). This is in accordance to the

findings of Sigana (2002) who while working on crab Thalamita crebata in Kenya

observed a similar trend in the class size of female 40.5-55.44 mm to be more numerous

than males. Males, however were reported being dominant in larger size classes ranging

from 55.4-80.44 mm.

Moreover, small growth rates of females is the result of higher energy allocation

for reproduction in females than males (Asakura 1992; Bertini and Fransozo 2000). In the

present study few juveniles (size <3cm CW) were reported, that too in the months of

March, August and September from the study area. The presence of juveniles during

spring and post rainy season viz March, August and September concides with the

abundance of phyto-zooplantons indicating nutrient accumulation during these months.

This is in accordance with the work of Litulo (2004) who found that the reproductive and

recruitment activity of fiddler crab Uca annulipes occurs in rainy season favouring food

supply and larval flux.


71

4.3.3: Breeding Season

Breeding season is the period when the female crabs carry eggs and perform

sexual activities. Presently as depicted in Tables 4.3.2 and 4.3.3, two peaks with the

breeding seasons viz June-July and December-January are evident with maximum

number of sexually mature male and female crabs. Of the total ovigerous females

collected, maximum (46.7%) belonged to the 4-5 cm size class (modal size class), 32.5%

in class size of 3-4 cm and remaining 18.3% in class size of 5-6 cm. This is in accordance

with the findings of Sudha Devi and Smija (2013) while working on crab Travancoriana

Schirnerae Bott (1696).

Our observations indicated that freshwater crab M.masoniana is a biannual

breeder witnessed by maximum adult population of male and females during peak

seasons and minimum/negligible number of juveniles in the month of March, August and

September from the study area. Such annual and seasonal breeding pattern has been

observed in semi terrestrial and intertidal brachyuran crabs (Sastri, 1983; Henmi and

Kaneto, 1989). The present finding are in line with study of Tongdee (2001) who

reported that in mud crab, Scylla species, two recruitments in a year occur indicating that

these mud crabs were seasonal breeder and breed twice in a year. A similar reproductive

pattern was evident in hermit crab D. brevirostris which displays seasonal reproduction

with several breeding peaks (Turra and Leite, 2000).


72

4.4: Studies on Morphometric parameters viz., size, sexual maturity, allometric

growth and length-weight relationship.

A total of 592 crabs, 282 males and 310 females were analyzed in the present

study. It was observed that there were more females than males. Sexual dimorphism was

quite conspicuous. Sex differentiating characters were observed in morphometric and

meristic features of both crabs. Male crabs have an inverted „T‟ shaped abdomen and

mature females have an „inverted U‟ shaped abdomen (Fig 8 a & b) Plate 8. M.

masoniana exhibit heterochely, one cheliped being larger than the other and this was

observed in both the sexes. The CW of males ranged from 2.0 to 6.3 cm, with chela

length (CHL) and chela depth (CHD) falling in the range of 1.0-5.7cm and 0.5-3.5 cm

respectively (Table 4.4.1). The mean body weight (MBDWt) of male M. masoniana

under study ranged from 16.674 to 90.186 gms (Table 4.4.2).

A look at the Table 4.4.1 exhibits males of M.masoniana in the range of 2.0 to 6.3

cm with respect to carapace width (CW). The chela length (CHL) range from 1.0 to 25.7

cm and Chela depth (CHD) ranged from 0.5 cm to 3.5 cm in the same species of male

crabs. The mean body weight (MBDWt) of the male crabs ranged from 16.674 to 90.186

gms. In females of the same species, the carapace width (CW) ranged from 2.0 to 6.0

with abdominal width ranging from 1.0-4.5cm. The mean body weight (MBDwt) in

females ranged from 18.7770-64.850gms.

The CW of females under study ranged from 2.0 to 6.0 cm with abdominal width

falling in the range of 1.0 to 4.5 cm (Table 4.4.1). The mean body weight (MBDWt) of

female M. masoniana under study was in the range of 18.7770 gms to 64.850 gms (Table

4.4.3).

The statistical analysis of the above data exhibited a positive allometry between

CW and all dimensions analyzed viz chela length, chela depth, mean body weight in

males; abdominal width and mean body weight in females. The allometric relations

showed different levels of growth, that were observed for the different values of the

coefficient b in each relation analyzed , but all the relations showed a positive allometric

growth. In males, the allometric growth exhibited significantly positive allometry in

relationship of CW V/s CHL and CW V/s Mean body weight (MBDWt) with values of


73

7.49 and 4.97 respectively whereas, in females, “slight positive allometry” was observed

in relation of CW V/s ABDW and CW V/s Mean body weight (MBDWt) with values of

3.55 and 1.07 respectively. (Table: 4.4.4)

Discussion

The present study identified the growth pattern of cheliped in case of males and

abdomen in case of females and body weight acting as the best indicators of

morphological sexual maturity. The relationship between chela size and carapace width

(CW) in the present study resulted in the morphological sexual maturity for males of M.

masoniana (Graph 4.4.4.1 and 4.4.4.2). The present observation, reports a positive

allometry of cheliped in adult individuals of M. masoniana which is consistent with the

predictions made by Hartnoll (1974), who compared the growth related to secondary

sexual characteristic in Brachyurans. In crab a pubertal molt is often associated with an

increase in cheliped size and allometric growth rate. Brachyuran males develop cheliped

for combat, display and courtship (Hartnoll, 1982). The significance of heterochely in

crab is unclear, but according to Daniels (2001) it may be related to sexual signaling and

defence and in females may indicate reproductive vigour as well as the ability to take

care off and protect their brood.

The present observation exhibited males with greater chela depth than that of the

females indicating larger cheliped in males as compared to females (Table 4.4.1). In

accordance to our findings, Akin Oriole et al., (2005) to have reported that the right

chelae diameter of males were significantly bigger (P< 0.05) than that of females in

C.armatum. On the similar lines other relevant parameters like body weight, chelae

diameter condition factor of male C. Pallidus were higher than those of females.

According to Mansur et al., (2005) in Dilocarcinus, pagei the positive allometry

found for the male cheliped propodus in the adult stage indicated the possibility of a

greater energy investment in the development of the structure after the prepubertal stage.

In the present study, carpace width (CW) was taken as independent variable for

studying the relationships between morphometrics and sexual activity. This is in

accordance with the findings of Aiken and Waddy (1989) who observed sexual difference


74

in the growth of several body parts relative to carapace size in lobsters. According to

Antekhai et al., (1994) carapace width is more appropriate index of body weight and an

ideal parameter for weight estimation. Present observation based on carapace width (CW)

as an independent variable is also in accordance with findings of Castiglioni and

Negreiros-Franzozo (2004) who reported that carapace width is the body dimension most

frequently used as an independent variable in the analysis of relative growth of crabs,

because it fully represent the physiological changes that occur through their life history.

During the present investigation it was found that the females reached

morphological sexual maturity at smaller size than the males. A look at the table 4.4.3

revealed that females of carapace width 2.5 cm showed isometric growth, whereas,

females of carapace width ranging from 3.5 to 4.5 cm showed positive allometric growth

and there after again isometric growth was exhibited at 5.5 cm carapace width in

females. Thus indicating 3.5 cm (CW) to be the size of morphological sexual maturity.

Males exhibited allometric growth at 4.5 cm indicating size of sexual maturity as

after this size cheliped showed isometric growth with respect to length and depth

measurements. The difference in the size at morphological sexual maturity between

males and females is consistent with the pattern proposed by Shine (1988) for

brachyurans. According to which, this pattern explained the requirement for reproduction

in two sexes. When females allocate their energy for reproductive purpose, such as

spawning and egg incubation, they tend to mature at smaller size than males, who invest

their resources in somatic growth and reach maturity at greater size.

Our findings are also in tune with the observations of Carsen et al., (1996) who

reported that sexual maturity occurred at 4.0 and 5.0 cm carapace width in males

Platyxanthus patogonicus and ovigerous females of same species who reach sexual

maturity between 42.7 and 72.0 mm of carapace width.

On similar pattern Silva and Chacur (2002) reported a morphological sexual

maturity of 13.3 mm carapace width for females and 14.4 carapace width for males for S.

rectum. Later Silva et al., (2007) however, reported sexual maturity in S. rectum with

carapace width of 17.4 mm for females and 18.5 mm carapace width for males. Recently


75

Castiglioni et al., (2011) reported morphological sexual maturity of 16.71 mm carapace

width for females and 15.73 mm carapace width for males in same species.

In the present study the positive allometric growth during the adult stage

correspond to the increase in the reproductive potential of the females in M.masoniana.

The characteristic of the adult stage has also been verified in other brachyuran species

(Pinheiro and Fransozo, 1993; Mantelatto and Fransozo, 1994; Negreiros-Fransozo et al.,

2003). The abdomen of females present an important reproductive function for most

freshwater crabs as they form a „chamber incubatory‟ with function of retaining the eggs

and newly hatched juveniles, as reported by Hines (1982). On similar lines our

observation on present study gets further authenticated, with respect to abdominal growth

as indicator of morphological sexual maturity (Graph 4.4.4.3).

Our observation further gets strengthened by the findings of Hartnoll (1974) who

reported that changes in allometric growth of female abdomen occur at the beginning of

sexual maturity. The inflection of the straight line in graphic analysis of ABDW/carapace

width clearly demonstrated the point at which sexual maturity is attained by female of

M. masoniana (Graph 4.4.4.3) indicating that the morphometric maturity was reached at

3.5 cm carapace width exhibiting positive allometry. Finney and Abele (1981) in

Trapezia frerruginea reported that abdomen growth rate decreases slightly as a result of

the sexual maturity with the fact that high positive allometric growth occurred in juvenile

females and isometric in adult females.

The present study also indicated positive allometry for relationship between

carapace width (CW) and mean body weight (MBDWt) in case of male and female M.

masoniana. A look at table 2 and 3 revealed that upto 4.5cm of carapace width females

were marginally heavier than males and thereafter males were heavier than females. A

significant positive allometry was indicated in males with values of 4.97 in males and

slight positive allometry with value of 1.07 in females (Table 4.4.4). (Graph 4.4.4.4 and

4.4.4.5).

Our findings are in accordance with the findings of Sukumaran and Neelkantan

(1997) who reported weight increase to be evident above a carapace width of 115 mm in

P. pelagicus one of the characteristic feature of present findings is the tendency of males


76

to be heavier than females and this observation matches well with those made by Potter

et al., (1983) in P. pelagicus and Prasad et al., (1989) in Scylla serrata.

The results of length weight relationship analysis in P.pelagicus indicated that in

juveniles and pre-adult crabs, weight gain is almost uniform, females being slightly

heavier than males until they attain 120-125 mm carapace width, thereafter males

becoming heavier than females at any given length (Josileen, 2011). In contrast to the

present findings, Dhawan et al., (1976) found the females of P. pelagicus to be heavier

than males at a given carapace width.


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4.5 Evaluation of nutritional status viz., Protein, Lipid, Ash and Moisture

content in the edible meat of M. masoniana and its relationship with

spawning activity.

The seasonal variations in the protein, lipid, ash and moisture content in female

crabs have been shown in Table 4.5.1

Protein

The mean average protein content recorded during the study period (Jan 2012-

Dec 2013) in the body meat of female M. Masoniana was 54.47± 0.42 with minima of

47.05± 0.83% in December i.e. during spawning period and maxima of 62.65± 0.35% in

March i.e. during post spawning period. (Table 4.5.1 and Graph 4.5.1). Protein showed a

highly significant and negative correlation with moisture, the values being r=-0.84311

and r=-0.86942 for Ist and IInd

year respectively. Values however, exhibit highly

significant and positive correlation with ash (r=0.659093 and r=0.649427) for Ist and IInd

year respectively. Proteins showed insignificant positive correlation with lipid

(r=0.549235 and r=-0.52197) for Ist and IInd year respectively.

Lipids

The mean average lipid content recorded in the body meat of female crab was

4.76± 0.42 with minima of 3.94± 0.27% in July i.e. during spawning period and maxima

of 5.75±0.45% in September i.e. post monsoon period. (Table 4.5.1 and Graph: 4.5.1).

Lipid showed an insignificant positive correlation with ash (r=0.452726 and r=0.469045)

in Ist and IInd year respectively. Lipid content, however, showed insignificant negative

correlation with moisture (r=-0.49769 and r=-0.45325) in 1st and 2

nd year respectively.

Ash Content

The mean average ash content was 8.39±0.44 with minima of 69±0.35 % in June

i.e. during spawning period and maxima of 12.29± 0.67% in March i.e. during post

spawning period. (Table 4.5.1 and Graph 4.5.1.). Again a rising trend was observed

during post monsoon period. There is no direct relationship between ash cycle and

spawning activity.


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Moisture

The mean average moisture content of female crab was 81.06±1.57 with minima

of 78.38±1.41% in April i.e. during post spawning period and maxima of 84.24±1.70% in

July i.e. during spawning period (Table 4.5.1 and Graph 4.5.1.). Minimum values were

evident during post spawning period (March and April) whereas increasing trend was

again witnessed during post monsoon period. Thus moisture exhibit an inverse trend

(relationship) with lipids and protein contents, while on insignificant and negative

correlation with ash (r=-0.99367 and r=-0.53252) in both years respectively could be

recorded.

Protein

In males proteins show higher percentage than females throughout the study

period .Males did not show any well marked seasonal fluctuation with respect to proteins

as they do not invest much energy for reproduction but use for somatic growth. As

evident from Table 4.5.2 and Graph 4.5.2 the mean average protein content recorded

during the study period in the body meat of male M.masoniana was 59.42±0.43 with

minima of 55.11±0.27% in October i.e. post monsoon period and maxima of

64.45±0.16% in April i.e. post spawning period. Protein recorded an insignificant and

negative correlation with moisture during both the years, the values being (r=-043661 and

r=-0.24393). Highly significant and positive correlation of ash with protein was another

characteristic observation during the present period of study with values being r=0.045

and r=0.1048 for two years respectively.

Lipids

A look at Table 4.5.2 and Graph 4.5.2 indicated the mean average lipid content in

the range of 5.17±0.47 with minima of 4.09±0.33% in December i.e. spawning period

and maxima of 6.56±0.42% in August i.e. post monsoon period. A study of table 4.5.1

and 4.5.2 further reveal that M.masoniana, males have higher lipid content than the

females and the seasonal variations in the lipid content were much more pronounced in

females as compared to males. Lipid showed an insignificant and negative correlation

with moisture, the value being r=-0.31735 and r=-0.43068 in 1st and 2



79

Ash

The mean average ash content recorded in the body meat of male crab was 8.58±

0.37 with minima of 6.69±0.49% in September i.e. post monsoon period and maxima of

11.48± 0.45% in June i.e. spawning period. (Table 4.5.2 and Graph 4.5.2).

Moisture

The moisture content in the male crab exhibited mean average value of

78.56±1.70 with minima of 76.16±1.51% in April i.e. post spawning period and maxima

of 80.82±1.82% in December i.e. spawning period. As evident from Table 4.5.1 and 4.5.2

male record less moisture content than females and show an inverse trend with proteins

and lipids. Moisture content was observed to be higher during spawning period (Dec-

July). Moisture content showed in significant and negative correlation with ash (r=-0.004

and r=-0.003) in 1st and 2


Discussion

The seasonal variations in biochemical composition of body meat of male and

female M.masoniana are presented in Table 4.5.1 and 4.5.2. Results are expressed on dry

tissue weight basis. Two year study revealed distinct variations in relation to varied

reproductive phases i.e., spawning period, post spawning period and post monsoon

period.

Proteins

Protein content in body meat of female crabs remained low during winters with

lowest average being 46.05% in the month of December and 48.51% in the month of July

during the two year study period. This lower percentage of protein during December-

July coincides with their spawning period when the gonads are in advance stage of

maturity. On the other hand, average protein values viz 62.16±0.30% and 63.15% were

recorded to be maximum during the month of March in two years of study period. This

higher percentage of protein content in female body meat during March coincides with

their non spawning period and a similar rise is also witnessed during post monsoon

period (Table 4.5.1). As evident from Table 4.5.1, protein content in female crab showed

marked seasonal fluctuations in the body meat. The pronounced fall in the protein content

in female tissues during spawning period (December and July) suggested mobilization of


80

protein from muscle to the gonads for their development. A similar trend has been

observed by Bakhtiyar (2008) in M. dayanum who recorded a fall in the muscle protein

which coincides with their spawning season when gonads were in advanced stage of

maturity. On similar lines Lambert and Dehnel (1974) reported that in crustaceans, a

great amount of energy gets channeled to the gonads during reproduction, which is

reflected in the deposition or depletion of nutrients with the advent or departure of

reproductive period. Our observations get authenticated by the work of Hislop (2001)

who recorded that during both wintering and spawning periods, food intake is

significantly reduced and fish depends entirely on stored energy resources for its survival.

A similar trend was observed by Sriraman (1978) in shrimp, Penaeus merguiensis and in

fresh water prawn M. idea.

High mean levels of proteins during post monsoon (October: 55.85%) and non

spawning phases (March: 63.15%) can definitely be attributed to planktonic blooms

during this period. Moreover, high levels of proteins during this period indicates the

diversion and deposition of proteins in the muscles, primarily aiming to meet the protein

requirement for the next breeding seasons. Our observations get strengthened by previous

recording made in the same stream where benthic species like chironomous larvae

(Tubifex sps) were abundant during summer and monsoon as indicated by work of

Sawhney (2004), Parvez (2005) and Nelofer (2005). Since the variations in protein is

influenced by feeding and breeding capabilities therefore, the protein cycle appears to

have a strong correlation with feeding and spawning phases which are further species

specific. Protein maxima and minima corresponding to the development/spawning and

biological regression/resting phases respectively, gets authenticated by work of Lee

(1986) on green mussel.

In males as evident from Table 4.5.2, the protein content did not show any well

marked seasonal fluctuation throughout the study period. It exhibited lowest value

55.10% in Oct (Post monsoon) and highest value of 64.50% in April viz. Post spawning

period and a stable range during spawning period. From the study it has been

hypothesized that males do not invest much energy for reproduction, but use most of it

for somatic growth i.e. males have lower energy requirement than females to form a fully

developed gonad (Kyomo, 1988; Jeckel, et al., 1989). When compared with males,


81

females had lower mean average protein content viz 59.42± 0.43 and 54.47± 0.42

respectively (Table 4.5.1 and 4.5.2).

Mean protein levels, therefore, were greater in male crab meat than females.

Along the same lines our findings further get authenticated by observations of Adeyeye

(2002) in freshwater crab like Sudanonautes africans and Sengul and Zeliha, (2011) in

Potamon potamios, who reported low protein values for female crab meat in comparison

to male. Our observations are in tune with the findings of Ozogul et al., (2010) who have

also reported variations in the protein contents of both male and female crab meat in

Callinectes sapidus. Present findings however, contradict to those reported by Khan

(1992) and Zafar et al., (2004) who found greater concentration of protein in female crab

meat than their male counter parts.

Crustacean muscles contain high concentrations of free amino acids such as

arginine, glycine, proline, glutamine and alanine (Cobb et al., 1975). Konsosu and

Yamaguchi (1982) have pointed out that the Free Amino Acids (FAA) content in

crustaceans is higher than those in fish and mollusc. Shell fish meat provides high quality

protein with all the dietary essential amino acids for the growth and maintenance of the

body (FNB, 2007). The nutritive value of any animal is decided by the presence of

essential amino acids. Total essential amino acid contributions are much higher in hard

shell crabs than the soft shell crabs. Free amino acids such as alanine, glutamine and

glycine are responsible for the unique flavor and taste of crab meat.

Lipids

Lipids are highly efficient source of energy and offer more than double the energy

released by carbohydrates and protein (Okuzumi and Fuji, 2000). Generally muscles of

Crabs and prawns contain lesser quantity of lipid (Bhavan, 2009) and fresh water crab

meat is generally low in fat as compared to marine crabs (Adeyeye, 2002).

As evident from Table 4.5.1, the lipid content in female crabs remain low during

monsoon (July) and winters (Dec-Jan) which coincides with their spawning season when

the gonads are in an advanced stage of maturity. The decline in the lipid content during

spawning period is possibly due to mobilization of lipid as energy source to meet the high

energy demands, during act of ovulation and spawning. Similar reports on energy


82

mobilization in fishes during spawning seasons have been made by Jafri and Khawaja

(1964), Love (1970) Kilne and Willet (2002) and Nargis (2006).

Wide fluctuations in muscle lipids have been reported to occur both in

hepatopancreas and gonads of prawns during gonadal development (Pillay and Nair,

1973). Similar trend was observed in M.dayanum by Samyal (2007) and Bakhtiyar (2008)

who recorded a fall in lipid content of muscles during spawning season when gonads are

in advanced stage of maturity.

It has been observed that in body meat the lipid content vary from 3.90 to 5.85%

in females and 4.08 to 6.60 in male crabs respectively. Our observations are in

accordance with the findings of Radhakrishnan and Natrajan (1979) who assessed the

lipid values in P. vigil from 5.13 to 9.73% those recorded by Radhakrishan (1979) in P.

pelagicus where the mean lipid value was 3.3 to 5.6%. On similar lines, Balasubramanian

and Suseelan (2001) too, reported the lipid values from 6.2 to 7.6% in C. smithi. A low

levels of lipid content have been observed in the body meat in mud crab (0.9-1.6%) by

Thirunavukkarasu (2005).

Presently, male crabs have been observed to have higher lipid content than female

crabs and such observations that male crabs necessarily have high lipid content is not a

universal phenomenon as contradictory results have been reported by Clarke (1980).

According to Clarke (1980) in many marine invertebrates, lipids are the most variable

fraction and low levels of lipids characterized males rather than females. Lipid content in

the present species is comparatively low as against higher values recorded in fin fishes.

Higher values of lipid content have been reported in I. crenata (5.4-15.6%) by Thomas

(1985) and P. vigil (16.8-31.9%) by Radhakrishnan (1979). Thus, the candidate species

has been placed in a better quality food list for human beings.

Freshwater crab meat is generally low in fat, which is good for health (Adeyeye,

2002). Generally, the muscles of crabs and prawns contained lesser quantity of lipid

(Bhavan, 2009). Therefore, crabs and prawns are preferred by the consumer. Freshwater

crab meat is low in cholesterol when compared to marine and brackish water species

(Sinha and Ahmed, 2011).


83

Ash

Ash content determines the total amount of micronutrient present in the tissue of

an animal. Micronutrients include mineral content present in the tissue. In the present

study , it has been observed that there is an increase in the level of ash content in the body

meat of M. masoniana (female) but male crabs, show no well marked seasonal

fluctuations in the ash content. As is evident from Table 4.5.1 and Graph 4.5.1 two peaks,

one in the month of March (12.35±0.72%) and second in the month of September (9.90 ±

0.83%) can clearly be recorded. This increase in ash content corresponds to the post

spawning period of female crab, and matches well with the recordings made by Love

(1970) who also have witnessed a similar rise in ash level during post spawning period.

Samyal (2007) and Bakhtiyar (2008) while investigating M. dayanum also

recorded two peaks in ash content in the months of February and August and have

attributed such variability in ash content to the utilization of mineral matter for the

growth and maturation of ovaries and spawning purposes. The mean average value of ash

in present study ranged from 8.39±0.44 to 8.58±0.37 in female and male crabs

respectively. (Table 4.5.1 and 4.5.2).

Our findings are in accordance with the findings of Adeyeye (2010) in

Sudananautes africans africans where ash concentration ranged between 4.60 to 14.92%

in whole body meat. On similar lines Gokoglu and Yerlikaya (2003) reported ash content

of blue crab Callinectus sapidus and swim crab Portums pelagicus in the range of

1.39±0.019 and 0.89±0.08% respectively for claw and body meat. As observed presently,

ash content in M.masoniana was higher (8.58±0.37%) than those reported in other fin

fishes and crustaceans viz M. rosenbergii (0.37%) by Gopakumar (1993); Catla Catla

(1.268%) and Labeo-rohita (1.021%) as reported by Jafri and Khawaja (1964). In

comparison to other fin fishes (1.09%-1.431% ) and other crustaceans (0.37%-22%) M.

masoniana could be a good source of micronutrients, with respect to high values of ash

content.

The Ash is left out after complete combustion of fish meat and gives a measure of

the total mineral content. The mineral serve as component of bones, soft tissues. The

calcium and phosphorus together account for 70 to 80% of the mineral in skeleton of fish


84

(Nair and Mathew, 2000). The hard shell crabs contributed maximum (3.985 mg) of

minerals (calcium>sodium>magnesium>potassium>zinc) and minimum was in soft shell

crabs (3.018mg) of mineral (calcium>magnesium>sodium>potassium>zinc). In both

cases calcium showed maximum concentration.

Moisture

Water as a component, contributed maximum to the chemical composition in the

muscle tissue of all organisms including fin fishes and shell fishes investigated so far.

The mean average value of moisture in present study ranged between 78.56±1.70 to

81.06±1.57 in male and female crabs respectively. (Table 4.5.1 and 4.5.2). That the

females have higher moisture content than males in the body tissue (body meat) is in

accordance to the findings of Kuley et al., 2008 who reported that male blue crab meat

had lower moisture content than female crab meat. Our result, however, are in

contradiction with the findings of Srinivasagam (1979) in S.serrata meat where moisture

content was found to be slightly higher in males than females. Similar observations were

made in the muscles of male prawns (Bhavan et al., 2010) were also higher levels of

water content have been recorded in males as compare to females.

In the present investigation, there is significant variation in the moisture content

of M. masoniana throughout the period of investigation. Comparatively high values of

moisture content were recorded during July and December viz 84.26±1.80% in females

and 80.98±1.89% in males. These two months viz July and December, happen to be the

breeding period of M. masoniana when they develop gonads and therefore a positive

correlation exists between moisture content and gonadal activity in the species under

observation.

Our findings are in accordance to the finding of Pillay and Nair (1970) and

Farragut (1965) who also reported high moisture content during breeding season in

Cancer magister. Similar fluctuation of moisture content in response to reproductive

cycle has been observed by Tagore (1990) during monsoon months as well as by Samyal

(2007) in M.dayanum. Samyal (2007) observed high moisture content during the months

of May-June and September-October which happen to be the breeding seasons of the

species as M. Dayanum is a bi-annual breeder. Bakhtiyar (2008) also observed similar


85

trend in M. Dayanum and L. rohita where the variation in the moisture content were

related with the spawning, related activities. The values for moisture content so obtained

in M masoniana are well within the range reported for other species of fin fishes and shell

fishes by Jafri and Khawaja (1964) viz. Tor putitora -77.98%; Labeo-rohita-78.37%

and Channa punctatus 76-43%. The moisture content ranged from 73.5 to 81.8% in body

meat of S. tranguebarica (Thirunavukkarasu 2005); 77.7% in the body meat of blue

swimming crab (Prasad and Neelkantan, 1989); 80.19% in S.serrata (George and

Gopakumar, 1987).

A point worth mentioning here is that increase in the moisture content

accompanied by a decrease in protein and lipid content in body tissue therefore moisture

exhibit an inverse relationship with lipid and protein contents. The higher percentage of

moisture noticed in body meat on one hand and the negative correlation observed

between water content and protein concentration of M.masoniana on the other hand are in

agreement with the observations made by Zafar et al., (2004) in S. serrata who observed

inverse relation might be due to low temperature, low feeding rate and high energy

demand to maintain body temperature and to cope up with food scarcity during winter.

Similar results highlighting the relationship between moisture and muscle protein have

been advocated by Nargis (2006).

These variations in the moisture content therefore might be due to spawning effect

and breakdown of lipid as a consequence of vigorous metabolism as supported by the

work of Idler and Bitners (1958) and Danbergs (1964). The average moisture content

recorded in the present study was higher than that reported for marine and intertidal crabs

(George and Gopakumar, 1987). According to Bassey et al., (2011), knowledge of the

moisture content of food stuff serves as a useful index of their keeping qualities and

susceptibility to fungal infection.


86

4.6 Macroscopic and Microscopic analysis of gonad development of M.

masoniana.

Seasonal variations in morphology of gonads of male and female crabs of M.

masoniana with respect to size and colour have been observed during the study period

(Jan. 2012-Dec. 2013). From Table 4.6.1, it is evident that carapace width of the

specimens ranged from 2.1-6.0 cm CW. The size at sexual maturity was found to be 3.5

cm CW for females and 4.5 cm CW for males. Mature individuals within the range of

mean size 3.5 cm (females) and 4.5 cm CW (males) were evident only during the

monthly collection of June-July and December-January.

Macroscopically, the extent of occupancy of body cavity by gonads as given in

Table 4.6.2, Fig 9 (a, b, c, d) and Table 4.6.3, Fig 11 (a, b) clearly reveled that

difference in gonadal stage varied in accordance with size colouration and volume

occupied in the body cavity (Plate 9 and 11). The macroscopic and microscopic

examination of the ovaries allowed the identification of four gonadal stages (Plates

9&10).

Stage- I (Immature)

Ovaries during this stage have been observed to occupy 1/6th

of the body cavity

and were transparent in appearance with no oocyte visible to naked eyes (Table 4.6.2.

Fig: 9). Histologically ovaries shows presence of rounded oogonia and follicular cells.

Oogonia are small oval cells enclosed in a thin lining of germinal epithelium undergoing

mitosis to form more oogonia or meiosis to form oocytes. (Fig: 10a).

Stage-II (ripening)

Ovaries occupied 1/4th

of the body cavity with white to yellowish in colour (Table

4.6.2 Fig: 9) Histologically some oocytes could be seen surrounded by follicular cells.

Primary oocytes are larger than oogonia with no longer mitotic division. Ooplasm is

characterized by absence of yolk. (Fig: 10b).

Stage–III (mature)

Ovaries occupied full body cavity and were orange red in colouration (Table

4.6.2, Fig: 9) Vitellogenesis begins with originating of yolk globules from perinuclear


87

yolk complex. Mature oocytes become enlarged further and were fully laden with yolk

vesicles. Follicular cells with nuclei surrounded the oocytes (Fig: 10c).

Stage-IV (spawned)

Ovaries occupy ¼ of the body cavity with pale yellow colouration and flaccid

appearance (Table 4.6.2 Fig: 9). Atresic oocytes showing atresia were observed. Atresic

stage is characterized by oocytes disintegration or fusion of nucleus with sinking of

ooplasm and collapsing oocyte membrane (Fig: 10d).

In males the testis exhibited varied colouration and consistency and were made up

of semniferous tubules (Table 4.6.3, Fig 11 and 12).

The macroscopic and microscopic examinations allowed the classification of the

testis in 3 stages: (Plates 11 & 12).

Stage I (immature)

The testis and vasa differentia occupy 1/6th

of body cavity and had transparent

coloration with gelatinus aspect (Table 4.6.3, Fig: 11a & b) Testicular follicles lined up

by a single layer of germinal epithelium giving rise to spermetogonial cells are observed.

Each primary spermatogonia under go mitotic division to form secondary spermatogonia

(Fig: 12a).

Stage: II (maturing)

The testis occupy 1/4th

of the body cavity and had creamy white colouration but

with gelatinous aspect. (Table 4.6.3, Fig: 11 a & b). Secondary spermetogonia

differentiate into spermatocytes by undergoing mitotic division but few resting

spermatogonia are observed forming new crop of germ cells for next breeding seasons.

(Fig: 12b).

Stage III (mature)

The testis occupy entire body cavity and had milky white colouration.

Spermatozoids were found in all seminiferous tubules (Table 4.6.3, Fig: 11 a & b).

Histological analysis of these stage indicated various cell types viz spermatogonia,

spermatocytes and large number of spermatids and spermatozoids in the lumen of

tubules. No germinal area were observed in semniferous tubules (Fig: 12c).


88

Discussion

The reproductive cycle in crustacean has been studied by many workers including

investigations on reproductive biology in relation to the monthly occurrence of mature

male and females and monthly distribution of individuals according to size, classes and

sex. (Colby and Fonseca 1984; Conde and Diaz 1989; Lopez-Greo et al., 2000; Moura

and Coelho 2000; Fransozo and Bertini, 2002). In the present investigations, based on

availability of data on carapace width (Table 4.6.1) it has been observed that

M.masoniana is a biannual breeder with the two breeding peak seasons viz June-July and

December-January. Maximum numbers of ovigerous females observed, found correlation

with maximum development of ovaries in the present studies on crab. Maximum

numbers of males with mature testis and maximum carapace width were observed in the

same seasons. The size at sexual maturity for female was 3.5 cm of CW and 4.5 cm for

males. These findings are in agreement with the work done by Sukamaran and

Neelakantan (1996) on relative growth and sexual maturity in the marine crabs Protonus

pelagicus.

In comparison to male crabs of the present study with size 4.5 cm , female crabs

are smaller viz., 3.5 cm of CW, which may be attributed to the reason that female crabs

spend large amount of energy in their gonadal cycle than male counterparts. Similar to

present observations Mantelatto (2003) also recorded differential changes in CW of male

and females indicating gonadal maturity in Mithraculus forceps of his studies. Our

findings are also in accordance with pattern proposed by Shine (1988) for brachyurans.

According to this pattern, the requirement for reproduction in two sexes. When females

allocate their energy for reproductive purpose, such as spawning and egg incubation, they

tend to mature at smaller size than males, who invest their resources in somatic growth

and reach maturity at greater size.

Observations on reproductive output per brood for brachyuran crab as advocated

by Hartnoll, (1985). To simply uphold present view point of strong correlation of body

size and weight within species. Our observations get further strengthened by work of

Mantelatto and Fransozo, (1997) on fecundity in crab Callinectes ornatus, who held that

the wide variability in carapace shape as well as abdomen width affects the volume

reserved for gonadal development. According to Conan et al., (2001) crustaceans may be


89

considered physiologically mature when they are capable of producing viable gametes.

Observations on presently studied crab reveled a series of morphological and

physiological events. (Table 4.6.1- 4.6.3 and Fig 9-12).

When external morphological characteristics of the gonads were compared to

histological descriptions, modifications that characterize the process in different

developmental stages of gonadal cells throughout the gonadal cycle could be clearly

demarcated. The macroscopic analysis of the present studies revealed that gonads witness

changes in volume as well as colour during the course of maturation (Table 4.6.2 and

4.6.3). It was observed that during maturation phases from stage I and IV the volume

occupied by maturing gonads in both male and female crabs ranged from 1/6th

of body

cavity to entire occupancy of the body cavity. This is in accordance to the findings of

Adiyodi and Subramonian (1983). Arculeo et al., (1995) who also held that during the

course of gonadal maturation the ovaries undergo a sequence of macroscopic changes in

its morphology viz relative size, which are easily delectable by naked eye.

The present study revealed that in females the colouration of the ovaries ranged

from transparent when immature (Stage I) to white in (Stage II) and yellow (Stage III)

and orange tones in (Stage IV). Present observations are in tune with the finding on

ovarian colour change during vitellogenesis as recorded for Cyrtograpsus angulatus by

Castiglioni santos (2001). Variations in colouration has also been used to elaborate

macroscopic scales in the ovarian development (Arculeo 1995 and Flores et al., 2002.)

In the present study too, the ovaries of M. masoniana showed a pronounced

macroscopic differentiation in size and colouration during the maturation process of the

gonad. During the vitellogenesis the amount of oocytes in secondary stage increases in

the ovaries as a result of yolk deposition resulting thereby a change in colouration from

yellow to orange red.

On similar pattern Charniaux-cotton (1980) observed that ovarian colouration is a

result of the storage of vitellogenine presenting carotenoid pigments. Goodwin (1951)

also held that change in colour is the result of modification in carotenoid content

occurring during oogeneis. The sequence in the change on colour of ovaries evidenced by

the accumulation of yolk has also been described for the fresh water crabs Eudaniela


90

garmani and Sinapotamon yangtsekiense by Rostant et al., (2008) and Chen et al., (1994)

respectively.

Histological analysis of female gonads of M. masoniana in the present studies

revealed a gradual process of oocyte development where cell types viz., oogonia, oocytes,

mature oocytes, atresic oocytes along with follicular cells were observed (Fig 10 a, b, c

and d). These observations indicating the description of cellular stages agrees with the

cell types observed in other female decapods crustaceans by Castiglioni et al., (2007);

Rostant et al., (2008), Souza and Silva (2009).

In this context, the presence of post ovulatory follicles in fresh water crabs as

reported by Rjeibi et al., (2010) and presence of these follicles responsible for

synchronic maturation ensuring the necessary amount of mature oocytes for spawning

gets authenticated by observations of Shinozaki-Mendes et al., (2011). Histological

analysis of the modification observed in the oocytes during the process of gonad

maturation as observed presently are similar to the description in the literature for other

females of decapods crustaceans (Adiyodi and Subramonian, (1983), Lopez et al., (1997),

Elorza and Dupre, (2000).

The general layout of reproductive system of M.masoniana was similar to those

found in other decapods (Cronin 1947; Ryan 1967; Joshi and Khana 1982; Castilho et al.,

2008). The system shows the bilateral symmetry and H shape, characteristic of many

brachyuran crabs. This arrangement is found in many crabs and crayfishes (Krol et al.,

1992; Cumberlidge 1999).

The morophological analysis of male gonads during course of maturation process

form stage I to III also indicated colour change from transparent to milky white (Table

4.6.3). Whereas, the histological analysis of these gonadal stages exhibited cell types viz.

spermatogonia, spermatocytes, spermatozoids which are responsible for the modification

of the coloration ranging from transparent in immature individuals to milky white in

mature ones (Fig: 12 a, b and c). These observations are in accordance with the findings

of Mota Alves (1975) and Castilho et al., (2008) who held that testicular lobules are filled

with large number of spermatocytes in different stages of development.


91

Seasonal changes in morphology and physiology in testies of M. masoniana is in

tune with findings of Joshi and Khanna (1982) who also reported similar changes in the

testis of fresh water crab Potamon koolooense, with different stages of development in

semniferous tubules. The changes in the colouration and cell types of male gonads of M.

masoniana under observation followed the pattern described for other decapods species

viz Ucides cordatus and Armases rubripes by Castilho et al., (2008) and Santos et al.,

(2009).

Chapter 5

Summary and Conclusion


92

Studies made on species diversity included its taxonomic position, morphological

characteristics, habits, habitat and distribution. Out of various water bodies

scanned so far two species of crabs were recorded viz Maydelliathelphusa

masoniana Henderson, (1893) from plain areas and Himalayapotamon

emphysetum Alcock (1909) from hilly stretches of the region. On the basis of

distribution pattern of two species viz Maydelliathelphusa masoniana and

Himalayapotamon emphysetum, it was observed that these two species reported

from the water bodies of Jammu region have contrast morphological features as

well as their distribution pattern. This variation in distribution pattern of the two

species clearly indicated the probability of presence of many more species in the

water bodies of different geographical conditions.

Physico-chemical parameters viz temp (Air/Water), pH, DO, FCO2, Cl-

, Ca++

,

Mg++

, HCO3 assessed during study period revealed abundance of crabs to be

more at station II as compared to station I as indicated by catch per unit effort


93

(C.P.U.E). It was observed that total crab population exhibited positive correlation

to water temperature (r=.009885), pH (r=0.338326), dissolved oxygen

(r=0.174293), calcium (r=0.056325) magnesium (r=0.602844), while negative

correlation was exhibited with chloride (r=-0.00447), HCO3-

(r=-0.245292) and

FCO2 (r=-0.396196). Based on the results of crab population viz.-a-viz.

physico-chemical parameters at two stations of Gho-manhasan stream of Jammu

region, J&K, it was concluded that water temp, depth pH, FCO2, DO, Ca2+

&

Mg2+

exhibit positive impact on crab abundance and therefore high C.P.U.E at

station II can be attributed to the combined effect of (a) favorable water quality

parameters (b) physical anthropogenic disturbances that are apparently very less

at station II.

Based on the relationship of carapace width (CW) with abdominal width

(ABDW) and mean body weight (MBDwt) in females and relationship of

carapace width (CW) with Chela length (CHL), Chela depth (CHD) and mean

body weight (MBDwt) in males, the size of sexual maturity and allometric growth

were assessed during the study period. A total of 592 crabs were obtained, 282

being males and 310 females. The maximum carapace width in female was 6.0

cm with abdominal width of 4.5 cm. Males exhibited maximum carapace width of

6.3 cm and chela length of 5.7 cm and chela depth of 3.5 cm. The mean body

weight in males varied from a minimum of 16.674 g to a maximum of 90.186 g.

Whereas in females, body weight ranged between 18.770 g being minimum &

64.850 g being maximum. Based on the data obtained from morphometric

analysis in male and female of M. masoniana the size at sexual maturity happened

to be 3.5 cm carapace width in females and 4.5 cm carapace width in males.

Knowledge of size frequency distribution, sex ratio and breeding season is of

great importance in the commercial utilization of a particular species. Various

parameters dealing with population structure of freshwater crab

Maydelliathelphusa masoniana, also incorporated their size frequency

distribution, sex ratio, and breeding season. A total of 592 crabs were obtained of

which 310 (52.36%) were females and 282 (47.63%) were males. The overall sex


94

ratio comes to be 1:1.09 with considerable degree of seasonal fluctuations. The

overall size frequency distribution exhibited males, females and juveniles. Sexual

dimorphism was characterized by the larger sized males in relation to females.

Breeding season indicated two peaks viz. (June-July & Dec-Jan), with scarce

young juveniles in the population.

The present investigation on the seasonal variation in the bio-chemical

composition of the body meat of M. masoniana, reveal that there are two distinct

period of variation i.e.

Spawning period ranging from June–July and December-January when the

gonads are in advance stage of maturity and there is mobilization of

nutrients from muscle to the gonads for their development.

Non-spawning period ranging from February to May (Post-spawning

period) and August to November (Post-monsoon period). During this

period M.masoniana is nutritionally more rich in protein, lipid, moisture

and ash content which are high in the muscles.

Further, it was observed that protein, lipid, ash and moisture content in

M.masoniana are comparable with other edible species of fin and shellfishes and

therefore qualify the criteria of edible food organism. The values however, are

considerably high during non spawning period. During this period macro nutrients

exhibit their maxima and therefore can be taken as the period of its harvesting and

consumption.

The morphological analysis of the gonads of both the sexes of M.masoniana when

compared to histological description exhibited modification during process in

different development stages throughout gonadal cycle. Gonadal stages of M.

masoniana exhibited that size ranging from 3.5-5.5 cm CW in case of both males

and females indicated the size at sexual maturity. The histological description was

based on 48 specimens (24 each sex). Four gonadal development stages were

found for females: immature, ripening, mature and spawned. Three development

stages were found for males immature, maturing and mature. Sexually mature


95

specimens (stage III) are found twice in a year (June, July and December-January)

exhibiting M. masoniana as biannual breeders. Morphological observation of

gonads at stage III exhibit (a) orange red colouration of ovaries in female (b)

milky white colouration of testis in males occupying full cephalothorax cavity.

Histological evidence further authenticated the gonadal maturity at stage III with

the presence of mature oocytes in ovaries and spermatozoids in testis.

Chapter 6

Recommendations

Recommendations

96

Crabs being highly sensitive to habitat loss, pollution and water siltation, their

disappearance qualify them as pollution indicators and can be intensively

investigated as possible bio-indicators of metals in sediments.

Rigurous survey need to be undertaken throughout the state as more species can

appear on record that will enrich the data on the existing crab resources of the

state.

Database on morphometry should be created on war footing as it can become

important document for the assessment of the crab fishery for their inclusion in a

regional food security programme by indicating minimum size of capture below

which harvesting should not be practiced.

Information on size frequency distribution, sex ratio and breeding season is very

important in formulating the strategy for initiation of crab culture.

Recommendations

97

Capture/Fishing of crabs during ovigerous/juvenile carrying females should be

banned and need to be declared as ‘no capture’/ ‘protected’ period on lines similar

to fishes.

Information on nutritional status can well be utilized to qualify M.masoniana as

an ideal item in dietary on similar lines like fishes as it possess high protein and

low lipid content and can serve a cheap source of animal protein.

Most important interaction between humans and crabs is later’s role as vector of

diseases. Crabs serve as secondary hosts of the diseases Paragonimiasis and

Onchocersiasis (river blindness). It is suggested that crabs should be consumed

after proper cooking only as raw crab meat can be a vector for above mentioned

diseases in human beings.

In rural areas, crabs which are rich in calcium are usually sought and eaten by

childless women in the belief that this would help them in attaining pregnancy.

This mythological aspect of crabs need to be studied and explored as it may then

help numerous childless couples which are so common during present times in

our society.

Smaller crabs which usually are not of great economic value can be used in the

preparation of high energy yielding cheaper artificial pelleted feeds which can be

used for cultivation of edible verities of crabs as well of aquaculture and poultry

feed.

The waste material of crabs viz; gills, intestine and shells is a good source of raw

material for high quality chitin and chitosan which can be of intense utility in

pharmaceutical and agriculture industries.

Many species of the crabs viz; Potamonautes (Malawi blue crab) of Lake Malawi

have been categorized as aquarium species. On similar corollary crab species

from Jammu waters can be searched and designated as aquarium wonders similar

to fishes.

Recommendations

98

Last but not the least, efforts should be done through advertisement so as to make

masses aware about crabs utility as an alternative source of nutritionally rich

animal protein for undernourished rural dwellers.

Thus, with such great importance and significance of lesser known ‘Crabs’ we

must ensure that enduring legacy is not to wipe out the small things that provide us with

great benefits ranging from culinary use to nutrient cycling and climate regulation.

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Appendix

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JOURNAL OF INTERNATIONAL ACADEMIC RESEARCH FOR MULTIDISCIPLINARY

Impact Factor 1.625, ISSN: 2320-5083, Volume 2, Issue 9, October 2014

146

Colour: All the specimens examined are characteristic in colour with dark

chocolate brown, shining carapace on dorsal side. The ventral surface of

carapace being light brown in colour.

Size: Species attains the size upto 7cm of carapace width (C.W), especially in

males. Whereas females of the same species are smaller with size upto 6 cm. of

carapace width (cw).

Heterochelous chelas: Heterochelous Chelas have been observed to be

present in both the sexes with heavy, unequal and bigger size chelipeds in

males than females. Cutting edges of chelas are armed with teeth of variable

size.All the teeth and tips of both fingers are dark brown in colour .

148

Colour: All the specimens examined have reddish orange to greyish brown

coloured dorsal surface, which is rough but not shining. The ventral surface of

carapace being pale white in colour.

Size: Specimens of this species are of moderate size, males reaching a size upto

5 cm of carapace width (cw). Females being smaller than males with size range

of 4.5 cm of carapace width (cw).

Heterochelous Chelas: Heterochelous chelas used for grasping defence and

sexual signaling are medium sized, less serrated but have a peculiar feature of

purple or blue with orange coloration at their tips.

studies on eco-biology of some freshwater crabs from jammu...

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