Biodiversity of demersal fish along the estuarine - shelf regions of Goa

Thesis submitted to Goa University

for the degree of Doctor of Philosophy

in Marine Sciences

53 7 By 7 7 D I

Vinay P. Padate

Department of Marine Sciences Goa University, Goa - 403 206, INDIA

October, 2010

5'2G Dedicated to

Late Shri (Dr) Kethavrao Yoshi Statement

As required by the University ordinance 0.19.8 (vi), I state that the present thesis entitled "Biodiversity of demersal fish along the estuarine — shelf regions of Goa" is my original contribution and the same has not been submitted on any previous occasion. To the best of my knowledge the present study is the first comprehensive work of its kind from the area mentioned. The literature related to the problem investigated has been cited. Due acknowledgements have been made wherever facilities and suggestions have been availed of.

Vinay P. Padate Certificate

This is to certify that the thesis entitled "Biodiversity of demersal

fish along the estuarine — shelf regions of Goa", submitted by Mr.

Vinay P. Padate for the award of Doctor of Philosophy in Marine Sciences is based on his original studies carried out by him under my supervision. The thesis or any part thereof has not been previously submitted for any degree or diploma in any Universities or

• Institutions.

Dr. C.U. • ivonker Research Guide Associate Professor Department of Marine Sciences Goa University Taleigao Plateau, 403 206, Goa

Ai I t•K& carr'eafrow) sw-jog*__ce .4-e-r6'e4 eczAAd)rbitP3 ACKNOWLEDGEMENTS

I would like to express a deep sense of gratitude to my guide, Dr. C.U.

Rivonker, for giving me an opportunity to work on a unique research problem under his able guidance.

I would like to express my gratitude to the Head, Department of Marine

Sciences and members of the FRC for their timely suggestions. Further, thanks are due to the Director, National Institute of Oceanography, Goa for allowing me to participate in the Ballast water Project which has been funded by the Directorate

General of Shipping, Government of India. I am highly indebted to Dr. A.C. Anil,

Senior Scientist, MCMRD, NIO and coordinator of the project for the timely discussions that enhanced my scientific aptitude. I would always be indebted to Dr.

S.S. Sawant, Senior Scientist, NIO for his help and support. I convey my gratitude to

Mr. K. Venkat, Senior Technical Officer, NIO for extending technical assistance, Dr.

Dattesh Desai, Scientist, NIO and Dr. Lidita Khandeparker, Scientist, NIO for their help in the DNA sequencing.

I am also indebted to Mr V. Khedekar, Senior Technical Officer, GOD, NIO for the Scanning Electron Microscope (SEM) photography of the crab specimens. The help extended by Dr. M.P. Tapaswi, Documentation Officer, NIO in obtaining the references from different places is gratefully acknowledged.

I also take this opportunity to express my deep gratitude to Dr. B.F. Chhapgar, an eminent carcinologist for his valuable suggestions in confirming the identity of the newly described species. Further, I acknowledge Dr. Mark Siddall, Curator at the

American Museum of Natural History, New York for his timely help in providing

Charybdis philippinensis specimens on loan for reference purpose. I thank the anonymous reviewers for the constructive criticism that enhanced the quality of the published papers.

I would whole — heartedly thank my colleagues Nutan and Tomchou for teaching me the intricacies of data handling, processing and presentation, Vineel for the artwork techniques, Mahabaleshwar and Ganesh for their help extended during my stay at the Department. A special thanks to my colleagues at the MCMRD, NIO

(Dhiraj, Rajat, Amar, Kirti, Lalita and Vinayak) for extending help during my work at the NIO. Finally, I take this opportunity to thank my close friends at the University,

Shivraj, Santosh, Sweety, Shilpa, Renosh and Milind for supporting me during the study period. I would also like to thank the non — teaching staff of the Department of

Marine Sciences for extending me all the help during my laboratory work. I wish to specially thank the fishing crew of the "Jesus Bless" trawler for allowing me to collect fish samples.

I wish to express deep sense of gratitude to Shri. Dada Samant, Smt. Suvarna

Teli, Shri. Prashant Padate and Shri. Satish Yedve for their blessings and support for my endeavour. I would also take this opportunity to specially thank Dr. Subodh

Sawant for his kind support and encouragement. I make a special mention of my mother Late Mrs. Vandana Padate, whose memory has been the sole source of inspiration on my path to seek knowledge and my parents, Mr. Pandharinath and Mrs.

Yamini Padate for their blessings, without which this endeavour would not have materialized.

Vinay Padate CONTENTS Page No. List of Tables i — ii List of Figures iii — vi

Chapter 1. General Introduction 1 1.1. Background information 1 1.2. Literature review 2 1.3. Objectives 6

Chapter 2. Materials and Methods 7 2.1. Study area 7 2.2. Sample collection 9 2.3. Taxonomic identification and morphometry 11 2.4. Sample preservation 12 2.5. 18S rDNA sequencing 12 2.6. Data compilation and processing 13 2.6.1. Faunal composition 13 2.6.2. Faunal abundance (numbers) and weight 13 2.6.3. Species diversity indices 14 2.6.3a. Shannon — Wiener's diversity index 14 2.6.3b. Heip's evenness index 15 2.6.3c. Margalefs species richness 15 2.6.3d. Simpson's dominance index 15 2.6.4. Cluster analysis 16 2.6.5. Functional guilds 16 2.6.6. Determination of catch rate 17

Chapter 3. Demersal faunal community structure off Goa 18 3.1. Introduction 18 3.2. Literature review 18 3.3. Results 22 3.3.1. Species composition 22 3.3.2. New records for the study area 23 3.3.3. Quantitative analysis 23 3.3.3 a. Overall data 23 3.3.3b. Crustaceans 26 3.3.3c. Fin fish 29 3.3.3d. Molluscs 36 3.3.3e. Other fauna 38 3.3.3f. Coastal residents and migrants 39 3.3.3g. Diversity 39 3.3.4. Faunal associations 41 3.3.4a. Family level 41 3.3.4b. Family level: pre — monsoon season 41 3.3.4c. Family level: post — monsoon season 42 3.3.4d. Species level 43 3.3.4e. Species level: pre — monsoon season 43 3.3.4f. Species level: post — monsoon season 44 3.3.5. Catch rates of the study area 45 3.4. Discussion 46 3.5. Conclusion 61

Chapter 4. Description of Charybdis (Charybdis) goaensis, new species 62 4.1. Introduction 62 4.2. Literature review 62 4.3. Taxonomy 65 4.3.1. Family Portunidae Rafinesque — Schmaltz, 1815 65 4.3.2. Subfamily Thalamitinae Paul'son, 1875 66 4.3.3. Genus Charybdis De Haan, 1833 66 4.3.4. Charybdis (Charybdis) goaensis sp. nov. 67 4.3.4a. Material examined 67 4.3.4b. Diagnosis 67 4.3.4c. Description 68 4.3.4d. Colour 71 4.3.4e. Etymology 71 4.3.4f. Distribution 71 4.3.4g. Habitat characteristics 71 4.3.4h. Habit 72 4.3.4i. Remarks 72 4.3.5. Charybdis (Charybdis) philippinensis Ward, 1941 74 4.3.5a. Original description by Ward (1941) 74 4.3.5b. Updated description of type specimens — C. philippinensis 76 4.3.6. Key to the species of swimming crabs of the sub — genus Charybdis 78 4.4. Conclusion 87

Chapter 5. A new record of Caesio cuning outside its known geographical array 88 5.1. Introduction 88 5.2. Literature review 88 5.3. Taxonomy 89 5.3.1. Family Caesionidae 89 5.3.2. Genus Caesio Lacepede, 1801 (Carpenter, 1988) 90 5.3.3. Caesio cuning (Bloch, 1791) . 90 5.3.3a. Material examined 90 5.3.3b. Description 91 5.4. Discussion 92 5.5. Conclusion 96

Chapter 6. A new record of Scylla olivacea from Goa — A comparative diagnosis 97 6.1. Introduction 97 6.2. Literature review 97 6.3. Taxonomy 102 6.3.1. Morph 1 103 6.3.1a. Material examined 103 6.3.1b. Description 103 6.3.2. Scylla olivacea (Herbst, 1796) description by Keenan et al. (1998) 105 6.3.3. Morph 2 105 6.3.3a. Material examined 105 6.3.3b. Description 106 6.3.4. Statistical analysis 108 6.3.5. DNA sequencing 108 6.3.6. Geographical distribution of Scylla olivacea 108 6.4. Conclusion 109

Chapter 7. Puffer fish diversity off Goa 110 7.1. Introduction 110 7.2. Literature review 110 7.3. Taxonomy 111 7.3.1. Family Tetraodontidae Fritzsche, 1982 111 7.3.2. Arothron immaculatus (Bloch & Schneider, 1801) 114 7.3.3. Chelonodon patoca (Hamilton, 1822) 115 7.3.4. Lagocephalus spadiceus (Richardson, 1845) 116 7.3.5. Takifugu oblongus (Bloch, 1786) 117 7.3.6. Genus Tetraodon Linnaeus, 1758 118 7.3.7. Tetraodon fluviatilis fluviatilis Hamilton, 1822 121 7.3.7a. Dekkers' (1975) description 121 7.3.7b. Present specimens — comparison with Dekkers' description 122 7.3.7c. DNA sequencing 123 7.4. Discussion 123 7.5. Conclusion 124

Chapter 8. Summary 125

Bibliography 127

Appendix List of Tables

Table 2.1. Details of trawl sampling carried out along the potential fishing grounds and bay — estuarine waters of Goa

Table 2.2. Details of shore sampling (beach seine and crab trap) carried out along the bay — estuarine waters of Goa

Table 2.3. List of meristic counts used in the identification of demersal marine fauna

Table 2.4. List of morphometric parameters used in the identification of demersal marine fauna

Table 2.5. Functional guilds used to categorize the demersal marine fauna along Goa coast

Table 3.1. List of demersal fauna identified during the present study from Goa, central west coast of India

Table 3.2. Size class and life stages of species observed during the present study

Table 3.3. New records of demersal marine species along Goa coast

Table 3.4a. Month wise differences in faunal abundance using two — way ANOVA (for months and faunal groups)

Table 3.4b. Season wise differences in faunal abundance using two — way ANOVA (for seasons and faunal groups)

Table 3.5a. Month wise differences in faunal weight using two — way ANOVA (for months and faunal groups)

Table 3.5b. Season wise differences in faunal weight using two — way ANOVA (for seasons and faunal groups)

Table 3.6. List of species categorized as "other teleosts" within the fin fish component of the demersal community during the present study

Table 3.7. Habitat use guilds of commonly observed demersal species during the present study

Table 3.8. Range and averages of species diversity indices computed during the present study

Table 3.9a. Month wise differences in diversity using one — way ANOVA (P = 0.001)

Table 3.9b. Month wise differences in evenness using one — way ANOVA (P = 0.001)

Table 3.9c. Month wise differences in species richness using one — way ANOVA (P = 0.001) Table 3.9d. Month wise differences in dominance using one — way ANOVA (P = 0.001)

Table 3.10a. Season wise differences in diversity using one — way ANOVA (P = 0.05)

Table 3.10b. Season wise differences in evenness using one — way ANOVA (P = 0.05)

Table 3.10c. Season wise differences in species richness using one — way ANOVA (P = 0.05)

Table 3.10d. Season wise differences in dominance using One — way ANOVA (P = 0.05)

Table 3.11. Ecological criteria observed for the major fish families during the present study

Table 3.12. Ecological functional guilds of the major fish families observed during the present study

Table 3.13. Catch rates of major fish groups during the present study

Table 3.14. Catch percentage of major fish groups off Calangute — comparison with Prabhu and Dhawan (1974)

Table 5.1. Details of morphometric measurements (in cm) of Caesio tuning specimens (N = 04) collected off Goa during the present study

Table 6.1. List of phenotypic characters and morphometric parameters (N = 24) used in morphometric analyses of mud crabs during the present study

Table 6.2. List of morphometric ratios (N = 27) derived for morphometric analysis of mud crabs during the present study

Table 6.3. Range, mean and standard deviation of morphometric ratios of S. serrata (N = 11) and S. olivacea (N = 20) obtained during the present study

Table 7.1. Morphometric measurements of marine puffers obtained during the present study along with their means and standard deviation

Table 7.2. Morphometric measurements of Tetraodon fluviatilis fluviatilis obtained during the present study (N = 04) along with their means and standard deviation

ii List of Figures

Fig. 2.1. Map of study area indicating coastal habitats, anthropogenic structures and sampling sites

Fig. 3.1. Taxa composition of demersal faunal groups observed during the present study

Fig. 3.2. Group wise composition of trawl catches observed during the present study (a) numbers and (b) weight

Fig. 3.3. Month wise variations in (a) total numbers and (b) total weight of major demersal faunal groups observed during the present study

Fig. 3.4. Month wise variations in (a) average numbers ± S.D. and (b) average weight ± S.D. of demersal fauna observed during the present study

Fig. 3.5. Composition of crustaceans in the trawl catches observed during the present study (a) numbers and (b) weight

Fig. 3.6. Month wise variations in (a) total numbers and (b) total weight of crustacean groups observed during the present study

Fig. 3.7. Month wise variations in (a) average numbers ± S.D. and (b) average weight ± S.D. of crustaceans observed during the present study

Fig. 3.8. Composition of fin fish in the trawl catches observed during the present study (a) numbers and (b) weight

Fig. 3.9. Month wise variations in (a) total numbers and (b) total weight of fin fish groups observed during the present study

Fig. 3.10. Month wise variations in (a) average numbers ± S.D. and (b) average weight ± S.D. of fin fish observed during the present study

Fig. 3.11. Composition of molluscs in the trawl catches observed during the present study (a) numbers and (b) weight

Fig. 3.12. Month wise variations in (a) total numbers and (b) total weight of mollusc groups observed during the present study

Fig. 3.13. Month wise variations in (a) average numbers ± S.D. and (b) average weight ± S.D. of molluscs observed during the present study

Fig. 3.14. Composition of other fauna in the trawl catches observed during the present study (a) numbers and (b) weight

Fig. 3.15. Month wise variations in (a) total numbers and (b) total weight of other fauna groups observed during the present study

iii Fig. 3.16. Month wise variations in average numbers ± S.D. and (b) average weight ± S.D. of other fauna observed during the present study

Fig. 3.17. Month wise trends in (a) average numbers ± S.D. and (b) average weight ± S.D. of coastal residents and migrant species observed during the present study

Fig. 3.18. Month wise variations in species diversity indices (average ± S.D.) of the demersal fish community during the present study

Fig. 3.19a. Dendrogram indicating similarity among fish families observed during the entire study

Fig. 3.19b. MDS plot indicating similarity among fish families observed during the entire study

Fig. 3.20a. Dendrogram indicating similarity among fish families observed during the pre — monsoon season

Fig. 3.20b. MDS plot indicating similarity among fish families observed during the pre — monsoon season

Fig. 3.21a. Dendrogram indicating similarity among fish families observed during the post — monsoon season

Fig. 3.2 lb. MDS plot indicating similarity among fish families observed during the post — monsoon season

Fig. 3.22a. Dendrogram indicating similarity among commonly occurring species observed during the entire study

Fig. 3.22b. MDS plot indicating similarity among commonly occurring species observed during the entire study

Fig. 3.23a. Dendrogram indicating similarity among commonly occurring species observed during the pre — monsoon season

Fig. 3.23b. MDS plot indicating similarity among commonly occurring species observed during the pre — monsoon season

Fig. 3.24a. Dendrogram indicating similarity among commonly occurring species observed during the post — monsoon season

Fig. 3.24b. MDS plot indicating similarity among commonly occurring species observed during the post — monsoon season

Fig. 3.25. Month wise trends in the weights of the jellyfish, Aurelia aurita and planktivorous fishes during the present study

Fig. 4.1. Generalized schematic diagram of the genus Charybdis indicating distinguishing morphological characters (a) Dorsal surface of carapace, (b) G1

iv Fig. 4.2. Charybdis (Charybdis) goaensis sp. nov. (a) Dorsal surface of carapace (Camera lucida diagram) (b) Dorsal surface of carapace (Photograph) (c) Frontal margin of carapace (Camera lucida diagram) (d) Antero — lateral margin of carapace (Camera lucida diagram) (e) Dorsal view of light cheliped (Camera lucida diagram) (f) Ventral surface indicating thoracic sternites and abdominal cavity of male (Camera lucida diagram) (g) Ventral surface indicating thoracic sternites and abdominal cavity of female (Camera lucida diagram) (h) Abdomen of male (Camera lucida diagram) (i) Abdomen of immature female (Camera lucida diagram) (j) Abdomen of mature female (Camera lucida diagram)

Fig. 4.3. Charybdis (Charybdis) goaensis sp. nov. Scanning Electron Micrographs (SEM) of First left pleopod or Gonopod (G1) (a) Entire Gl, (b) Tip of Gl, (c) Tip of G1 (enlarged view)

Fig. 4.4. Charybdis (Charybdis) philippinensis Ward, 1941 (a) Dorsal surface of carapace (Photograph) (b) Ventral surface of carapace (Photograph) (c) Frontal margin of carapace (Camera lucida diagram) (d) Antero — lateral margin of carapace (Camera lucida diagram)

Fig. 5.1a. Distinguishing characters and colour of fresh specimen of Caesio cuning (Photograph)

Fig. 5.1b. Caudal fin of Caesio cuning indicating characteristic yellow colouration (Photograph)

Fig. 5.1c. Ventral portion of Caesio cuning indicating characteristic pink colouration (Photograph)

Fig. 5.2. Supra — temporal band of scales in Caesio cuning (a) Photograph, (b) Diagrammatic representation (Carpenter, 1988)

Fig. 5.3. Dorsal fin of Caesio cuning indicating scaled portion and characteristic fin count (Photograph)

Fig. 5.4. Anal fin of Caesio cuning indicating characteristic fin count (Photograph)

Fig. 5.5. Map indicating the worldwide distribution of Caesio cuning

Fig. 5.6. Seasonal variations in hydrographic parameters measured at surface and 10 m depth during 2007

Fig. 6.1. Scylla serrata (a) Dorsal view of carapace (Photograph), (b) Frontal view of carapace (Photograph)

Fig. 6.2. Scylla serrata — Frontal margin of carapace (Camera lucida diagram) Fig. 6.3. Scylla serrata — Right cheliped (Photograph)

Fig. 6.4. Scylla serrata — Colour pattern on female abdomen (Photograph)

Fig. 6.5. Scylla serrata (a) Entire Gl, (b) Tip of Gl, (c) Tip of 01 (enlarged view) (Camera lucida diagrams)

Fig. 6.6. Scylla olivacea (a) Dorsal view of carapace (Photograph), (b) Frontal view of carapace (Photograph)

Fig. 6.7. Scylla olivacea — Frontal margin of carapace (Camera lucida diagram)

Fig. 6.8. Scylla olivacea — Right cheliped (Photograph)

Fig. 6.9. Scylla olivacea — Colour pattern on female abdomen (Photograph)

Fig. 6.10. Scylla olivacea (a) Entire G1, (b) Tip of G1, (c) Tip of 01 (enlarged view) (Camera lucida diagrams)

Fig. 6.11. DNA sequence of Scylla olivacea obtained during the present study (a) Aligned sequence data (385 bp), Alignment view and Distance matrix table, (b) Phylogenetic tree indicating position of the present sample

Fig. 6.12. Map indicating the worldwide distribution of Scylla olivacea

Fig. 7.1. Marine puffer fish species obtained during the present study (a) Arothron immaculatus, (b) Chelonodon patoca, (c) Lagocephalus spadiceus, (d) .Talcifugu oblongus (Photographs) '

Fig. 7.2. Diagrammatic representation of colour pattern on (a) sides and (b) back of Tetraodon fluviatilis fluviatilis (redrawn and modified from Dekkers, 1975)

Fig. 7.3. Colour pattern on the sides of Tetraodon fluviatilis fluviatilis collected during the present study (a) Photograph (b) Diagrammatic representation

Fig. 7.4. Colour pattern on the back of Tetraodon fluviatilis fluviatilis collected during the present study (a) Photograph (B) Diagrammatic representation

Fig. 7.5. DNA sequence of Tetraodon nigroviridis obtained during the present study (a) Aligned sequence data (584 bp), Alignment view and Distance matrix table, (b) Phylogenetic tree indicating position of the present sample

vi Chapter 1. General Introduction 1.1. Background information

Biodiversity means "the variability among living organisms from all sources including inter — alia, terrestrial, marine and other aquatic systems and the ecological complexes of which they are part; this includes diversity within species, between species and of ecosystems" (UNEP, 1992). A great deal of effort has been made to address issues pertaining to terrestrial and freshwater biota and their inherent ecological processes. However, the marine environment, although occupying more than 70 % of the earth's surface, has received little attention in spite of the fact that thirty five known phyla inhabit the marine ecosystems and fourteen among these are endemic (Sala and Knowlton, 2006). It is only in recent times, advances in sampling technology enabled to unravel the genuine scenario of marine biodiversity.

The demersal environments form a complex mosaic of bottom habitats and microniches those support more than 90 % of the total species. Further, these environments are dynamic with regard to the major ecological processes such as riverine influx, nutrient cycling and regeneration those sustain biomass and yield approximately 40 % of the global fish catch (Grainger and Garcia, 1996). On the other hand, the coastal waters are vulnerable to diverse phenomena resulting directly or indirectly from various anthropogenic activities as envisaged by the demographic pressure imposed by approximately 60 % of the total world population residing in

coastal areas through problems stemming from sewage discharge, over — fishing and

land use pattern (Taylor Jarnagin, 2004). Another significant factor that has adverse

effects on the demersal environment is bottom trawling, an indiscriminate non —

selective gear that removes millions of tonnes of non — targeted species resulting in

loss of diversity through habitat destruction (Norse and Watling, 1999). This

phenomenon over long term causes a shift in the marine food web having serious

1 implications for exploitable stocks. Recently, large — scale shipping activities are also considered to threaten the native biota through introduction of harmful invasive species altering the ecosystem function, thus affecting fisheries (Anil et al., 2002).

Goa, along the central west coast of India represents a paradigm of the ill — effects of unrestrained and inconsiderate human development on the environment

(Noronha, 2010). An integrated effect of these activities has put tremendous pressure on coastal ecosystems those affect the fishery leading to decline in the overall catch per unit effort (CPUE), and resources such as prawns have been over — exploited

(Ansari et al., 2006). Secondly, rapid rate of urbanization, industrialization, mining and shipping along its coastal region are potentially hazardous to its inland waterways and adjacent coastal waters with undesirable effects on coastal productivity.

The above mentioned imminent threats to the sensitive coastal ecosystems necessitate an appropriate strategy for management and conservation of living resources through sustainable use enabled by the development of risk assessment practices. The primary step in this regard involves development of database of the available resources through the use of conventional taxonomy and molecular techniques. Further, a comprehensive assessment of the environmental variables and coastal processes those bring about the coupling between pelagic and demersal environments is essential to discern the critical pathways associated with trophic relations those play an important role in the ecosystem function.

1.2. Literature review

The history of inventorying fauna dates back to the fourth century BC, when

Aristotle (384 — 322 BC) identified and classified fauna into groups such as insecta, crustacea and testacea (molluscs). The recognition of this was spearheaded by Carl

2 von Linne (or Carolus Linnaeus) through his classical work "Systema Naturae"

(1758). However, the research on marine fauna received impetus through several major maritime research expeditions (HMS Beagle, HMS Challenger, HMS

Endeavour) and voluminous works by some of multitude of naturalists (Forskal,

Fabricius, Lacepede, Cuvier, Lamarck and Bloch) during the eighteenth and nineteenth centuries. The subject was revolutionized by Haeckel (1866) through establishment of the "Phylogeny" concept, which was put into practice only during the twentieth century, wherein anatomy, chromosomes, biochemistry and protein analysis were used in concordance with phenetics. Further, Hennig (1966) proposed a new technique called "Cladistics", wherein similarities those grouping species were used in classification, and hypothesis related to systematics employed the rule of monophyly.

Most of the above efforts were appropriately supported by the establishment of natural history museums (Cato and Jones, 1991), gene banks (Benson et al., 2008) and electronic databases (Encyclopedia of Life, 2010; Froese and Pauly, 2010; Global

Biodiversity Information Facility, 2010; Palomares and Pauly, 2010) through preservation of biological specimens, genetic material, published literature and taxonomic databases.

Bloch (1785, 1787, 1801) pioneered the studies pertaining to marine fauna of the Indian region with special emphasis on fishes. Subsequently, several efforts were made during the eighteenth and nineteenth centuries to describe fishes from the southern and eastern coasts of India (Lacepede, 1803; Russell, 1803; Hamilton, 1822;

Bleeker, 1853; Blyth, 1858, 1860a, 1860b; Day, 1865). However, the first comprehensive attempt to compile information on marine fishes of the Indian coasts was made by Sir Francis Day (1876 — 1878, 1888, 1889), wherein detailed accounts

3 of approximately eleven hundred marine and estuarine species from the Indian sub — continent were documented.

Alfred William Alcock carried out the most prolific taxonomic work on wide array of marine fauna collected on — board the "RIMS Investigator" from the seas off

India and adjoining British colonies. His publication entitled "The Carcinological

Fauna of India" (1895, 1896, 1898a, 1899a, 1899b, 1900) contains descriptions of six hundred and five marine brachyura including those of one hundred and twenty six new species. In addition, he prepared descriptive catalogues of twenty five deep — sea madreporarian corals (1898b), one hundred and sixty nine deep — sea fishes (1899c), twenty seven dromidean brachyura (1901a), one hundred and seventeen decapod crustacea — macrura and anomala (1901b), eighty nine anomura (1905), twenty one penaeid prawns (1906) housed in the Indian Museum, Kolkata. Gardiner (1903 —

1906) compiled two volumes on the marine fauna of Lakshadweep and Maldives archipelagoes. Kemp (1915) provided detailed descriptions of the marine fauna of the

Chilka Lake: Thereafter, efforts were also made during the post — independence period (Silas et al., 1983; Mookherjee, 1985; Kurian and Sebastian, 1986; Rao and

Rao, 1993; Apte, 1998; Rao, 2003; Raje et al., 2007) to create nationwide inventories of various marine faunal groups.

In addition, several institutions such as the Royal Asiatic Society of Bengal,

Indian Museum (Kolkata) and Bombay Natural History Society (Mumbai) strived to uphold the cause of marine diversity documentation through taxonomic expertise and museum facilities.

A review of regional level studies reveals voluminous amount of

documentation from the eastern coast of India by the Zoological Survey of India through its "State Fauna Series" (Rao et al., 1991, 1992) and other occasional

4 publications (Deb, 1995; Rao et al., 2000; Mishra and Krishnan, 2003; Barman et al.,

2007). The Centre of Advanced Study in Marine Biology at Parangipettai has contributed significantly to the marine database documentation along the southeast coast of India (Sethuramalingam and Khan, 1991; Kasinathan et al., 1997; Rajagopal et al., 1998; Ramaiyan and Senthil Kumar, 1998; Jeyabaskaran et al., 2000). In addition, several others (Goswami, 1992; Sujatha, 2005; Ramesh et al., 2008) have contributed to the documentation of marine fauna of various coastal states. Much of the research pertaining to the Indian marine fisheries focused primarily towards reporting the commercial species and tertiary production potential of the coastal waters (Rao and Dorairaj, 1968; Bapat et al., 1972; Prabhu and Dhawan, 1974;

Radhakrishnan, 1974; Rao, 1988; Mathai et al., 1998; Rajkumar et al., 2005).

Studies pertaining to the diversity of marine demersal resources from the estuarine and shelf waters of Goa are scanty and do not provide systematic information on the biogeography of these waters. Published reports (Tilak, 1973;

Talwar, 1973) based on the occurrence of finned fishes of Goa suggest that the coastal waters of Goa harbour diverse pelagic and demersal fin fish assemblages. These reports are based on the collection of biological specimens thirty to fifty years prior to their publication, thus provide a preliminary insight into the species composition.

Ansari et al. (1995, 2003) attempted to describe the ichthyofaunal community structure from the bay — estuarine waters of Goa based on the seasonal variations in their occurrence, distribution, species diversity and attempted to elucidate the role of environmental factors on the trawl catches of this region, respectively. In addition to these, very few attempts aimed at providing preliminary information on the taxonomic status of various macro — fauna of some of the selected habitats in the region (George,

1980; Parulekar et al., 1980; Chatterji, 1994; Lobo, 2005). However, little efforts

5 were being made to elucidate different components of the demersal fish assemblages those include both rare and commercial species.

In view of the above, the creation of a comprehensive database on the demersal marine fauna was pertinent to provide a platform towards improved understanding of the coastal diversity of Goa. The present study primarily attempts to provide baseline information on the species composition of demersal fauna to facilitate creation of an inventory encompassing all the components of the demersal community. Secondly, the study attempts to describe the demersal community structure inclusive of faunal abundance, diversity, species associations and to evaluate the fishery potential of the region through quantitative analyses of trawl data. Lastly, molecular techniques have also been used to determine genetic variability among few newly reported species.

1.3. Objectives

• To study the demersal fish composition of trawl catches in the shelf waters of

Goa, central west coast of India.

• To quantify and identify the dominant species and assess their role in

ecosystem function.

• To determine genetic variability among newly reported species to assess their

invasive nature.

6 Chapter 2. Materials and Methods 2.1. Study area

Goa, with a coastline of about 105 km along NNW — SSE, facing the Arabian sea supports diversified geological and ecological features and forms an integral part of the central west coast of India (Wagle, 1993). The seabed consists of silty — clay up to 50 m and sandy — silt from 50 to 100 m (Modassir and Sivadas, 2003) with an average slope of 1.50 m.km-I up to approximately 55 m depth, and the submarine contours are approximately parallel to the coastline (Veerayya, 1972). The bathymetry is intermittently interrupted by coral reefs (Rodrigues et al., 1998) and submerged rocky patches those extend from the cliffs and promontories along the adjacent rocky shores (Wagle and Kunte, 1999). The overlying waters perennially receive nutrient — rich freshwater influx from the adjoining estuaries, particularly the Mandovi — Zuari estuarine complex (between 15°25'N and 15°31'N and between 73°45'E and

73°59'E) being the most prominent with catchment area of 1700 km 2 (Qasim, 2003).

The two major rivers namely Mandovi and Zuari are connected to the Arabian sea by the Aguada and Mormugao bays, respectively. The Aguada bay (4 km long) extends in north — south direction (Shetye et al., 2007); the Mormugao bay (14 km long) extends in an east — west direction from the Western Ghats, and the rock outcrops extending in a north — south line across the entrance of the bay separate it from the

Arabian sea (Rao and Rao, 1974). The tides are of semi — diurnal nature (Qasim and

Sen Gupta, 1981) and carry the seawater up to a considerable distance upstream.

The above region experiences maximum precipitation during the southwest monsoon accompanied by stormy weather, while quieter conditions prevail during the rest of the year (Ansari et al., 1995). The intertidal estuarine marshy ecosystem is the transformation of gentle sloping of nearshore banks of Mandovi and Zuari, which is filled with silt, clay and detritus transported by riverine influx from upper reaches,

7 where mangrove vegetation occurs in high density. The marshy areas extend for a distance of 4 km and inundated during high tide. The entire mudflats consist of loose muddy soil bordered by mangrove vegetation, thus making it highly productive for benthos those support large number of economically important species (Ansari et al.,

1995; Kulkarni et al., 2003).

However, in recent years the region has been subjected to large — scale developmental activities, those perpetuate anthropogenic pressure due to mining in the eastern part of its catchment area and the ore — transport along its riverine channels (Nigam et al., 2002), aquaculture farms (De Sousa, 2007), disposal of sewage (Ramaiah et al., 2007) and agricultural effluents (Sardessai and Sundar,

2007). All the above activities release wastes into the estuarine environment thereby altering the water quality and endangering the estuarine biota. Moreover, the shipping and dredging activities at the Mormugao Port (15°25'N, 73°47'E) located at the western end of the Mormugao bay (Rao and Rao, 1974) are also potential threats to the benthic habitat structure and the marine food web, with probable consequences on the community (Allan et al., 2008).

The adjacent coastal waters constitute the potential fishing grounds off Goa

coast and are subjected to intensive fishing by traditional and mechanized fishing crafts those employ variety of fishing gear (trawl net, purse seines, gill nets) to exploit the abundant pelagic and demersal resources (Rao and Dorairaj, 1968; Prabhu and

Dhawan, 1974; Ansari et al., 1995, 2003) with the exception of the seventy five — day

legislative ban on fishing (Goa, Daman & Diu Marine Fisheries Rules, 1981). The mechanization of fishing crafts, particularly the bottom trawlers and the subsequent

expansion in the fishing activity has led to intensive exploitation of the fish resources.

Intensive long — term trawling activity may lead to habitat destruction (Thrush and

8 Dayton, 2002), reduction in community complexity (Collie et al., 2000) as well as alteration in the composition and • structure of the resident faunal assemblages

(Longhurst and Pauly, 1987). Further, the adjacent nearshore coastal ecosystems are vulnerable to accidental oil spills (Fondekar, 2005) and shipwrecks. In 2000, an ore — carrier, MV River Princess was grounded off Candolim — Sinquerim (North Goa) due to inclement weather conditions (Ingole et al., 2006). The ship's contents spilled onto the neighbouring sandy shore leading to adverse consequences on the native benthic faunal assemblages. However, the grounded structure may have created an artificial reef — like habitat (Padate et al., 2010a), thus facilitating recruitment of juveniles of reef inhabiting fishes (Arena et al., 2007).

The present study area (Fig. 2.1) encompasses the nearshore fishing grounds located between 15°32'N and 15°28'N and between 73°45'E and 73°48'E (up to 25 m depth) off the Baga estuary — Aguada bay region, lower regions of the Mandovi —

Zuari estuarine complex including the Cumbharjua canal and the adjacent bays.

2.2. Sample collection

The present study encompassed four years of faunistic surveys along the bay — estuarine and nearshore coastal waters of Goa, central west coast of India, up to 25 m depth (Fig. 2.1) to assess the diversity and community structure of the demersal fauna.

The sampling surveys comprised bottom trawls, beach seines and crab traps with a total sampling effort of one hundred and sixty hours. The core of the sampling comprised bottom trawls (N = 95) with a total effort of one hundred and fifty six hours (Table 2.1). Among these, nineteen trawl hauls were taken from the bay — estuarine waters during May and December, 2005, September — October, 2006,

February, May, September and December, 2007 and January, 2008 (six in Mormugao

9

73°35E 73°40'E 73°45'E 73°50'E 73°55'E

15°30.N

15(75.N

15°20'N ON=

4411 ROCKY SUBSTRATUM II CLAYEY SUBSTRATUM 1 SANDY SUBSTRATUM SILTY-CLAY SUBSTRATUM 11 MANGROVE POTENTIAL FISHING COPALREEF PORT VEGETATION I 1 GROUNDS 1T1 POSITION OF BEACH SEINE CHANNEL CRAB TRAP 1=1 ICI /A V RIVER PRINCESS I I OPERATION el DEPLOYMENT GILL NET SITE OF COLLECTION TRAWL OPERATION SITE OF COLLECTION 1 0 1 COLLECTION 1 I OF C. gotensia [ c I OF C. curving •

Fig. 2.1. Map of study area indicating coastal habitats, anthropogenic structures and sampling sites Table 2.1. Details of trawl sampling carried out along the potential fishing grounds and bay - estuarine waters of Goa

Sr. Sampling Sampling area Geographical Position Sampling Sampling No. Date depth (m) duration (min) 1. 12.05.2005 Mormugao bay 15*24'37.7"N, 73*48'43.4"E 05 - 06 60 to 15°24'38.8"N, 73°50'07.1"E 2. 12.05.2005 Mormugao bay 15°26'48.3"N, 73°48'27.4"E 05 - 06 60 to 15°25'59.2"N, 73°50'40.6"E 3. 13.12.2005 Mormugao bay 15°24'41.2"N, 73°48'56.6"E 05 - 06 60 to 15°24'41.9"N, 73°51'11.0"E 4. 21.01.2006 PFG* 15°28'54.0"N, 73°45'18.0"E 05 - 08 76- to 15°29'24.0"N, 73°43'30.0"E 5. 21.01.2006 PFG* 15°30'06.0"N, 73°44'18.0"E 06 - 07 85 to 15°31'30.0"N, 73°43'06.0"E 6. 21.01.2006 PFG* 15°32'30.0"N, 73°42'42.0"E 07 - 09 72 to 15°32'12.0"N, 73°42'18.0"E 7. 21.01.2006 PFG* 15°32'48.0"N, 73°44'06.0"E 05 - 06 98 to 15°31'24.0"N, 73°44'18.0"E 8. 21.01.2006 PFG* 15°29'24.0"N, 73°44'54.0"E 03 - 05 80 to 15°29'06.0"N, 73°47'12.0"E 9. 25.02.2006 PFG* 15°31'30.0"N, 73°41'12.0"E 1 c - 21 110 to 15°32'54.0"N, 73°40'06.0"E 10. 25.02.2006 PFG* 15°32'30.0"N, 73°40'30.0"E 18 - 20 105 to 15°31'12.0"N, 73°41'18.0"E • 11. 25.02.2006 PFG* 15°31'48.0"N, 73°41'48.0"E 17 - 18 95 to 15°32'42.0"N, 73°41'24.0"E 12. 25.02.2006 PFG* 15°32'48.0"N, 73°41'48.0"E 16 - 19 100 to 15°31'30.0"N, 73°41'24.0"E 13. 27.03.2006 PFG* 15°30'12.0"N, 73°45'48.0"E 05 - 07 120 to 15°31'06.0"N, 73°45'18.0"E 14. 27.03.2006 PFG* 15°30'48.0"N, 73°45'12.0"E 09 - 13 155 to 15°29'54.0"N, 73°44'54.0"E 15. 27.03.2006 PFG* 15°29'48.0"N, 73°44'24.0"E 12 - 14 175 to 15°32'48.0"N, 73°43'12.0"E 16. 24.04.2006 PFG* 15°30'06.0"N, 73°41'48.0"E 15 - 22 71 to 15°32'30.0"N, 73°41'18.0"E 17. 24.04.2006 PFG* 15°32'48.0"N, 73°40'42.0"E 1, - 22 86 to 15°30'06.0"N, 73°41'42.0"E 18. 24.04.2006 PFG* 15°29'54.0"N, 73°42'18.0"E 18 - 20 107 to 15°32'12.0"N, 73°41'48.0"E 19. 24.04.2006 PFG* 15°32'06.0"N, 73°41'12.0"E 18 - 20 79 to 15°30'42.0"N, 73°41'54.0"E 20. 29.09.2006 Mormugao bay 15°24'41.2"N, 73°48'56.6"E 06 - 07 60 to 15°24'41.9"N, 73°51'11.0"E 21. 6.10.2006 PFG* 15°29'37.2"N, 73°45'19.7"E 10 - 12 60 to 15°31'25.7"N, 73°43'57.6"E 22. 6.10.2006 Mormugao bay 15°24'37.7"N, 73°48'43.4"E 05 - 06 60 to 15°24'38.8"N, 73°50'07.1"E 23. 6.10.2006 Zuari estuary 15°25'54.0"N, 73°50'51.0"E 05 - 06 60 to 15°26'38.5"N, 73°49'25.4"E 24. 9.12.2006 PFG* 15°29'41.2"N, 73°43'56.6"E 14 -16 192 to 15°32'41.9"N, 73°42'11.0"E 25. 9.12.2006 PFG* 15°32'41.9"N, 73°42'11.0"E 14 - 16 188 to 15°29'41.2"N, 73°43'56.6"E 26. 29.12.2006 PFG* 15°30'05.6"N, 73°43'25.0"E 14 - 16 125 to 15°33'47.7"N, 73°42'23.6"E 27. 29.12.2006 PFG* 15°33'42.0"N, 73°42'15.8"E 14 - 16 150 to 15°29'55.3"N, 73°43'27.7"E

28. 29.12.2006 PFG* 15°30'01.5"N, 73°43'24.1"E 0 ' ' 15 125 to 15°30'09.4"N, 73°45'12.7"E 29. 6.01.2007 PFG* 15°29'32.6"N, 73°45'19.0"E 05 -09 108 to 15°32'25.1"N, 73°45'07.7"E 30. 6.01.2007 PFG* 15°32'23.4"N, 73°45'07.0"E 06 - 09 129 to 15°32'23.5"N, 73°45'10.1"E 31. 6.01.2007 PFG* 15°32'21.0"N, 73°45'08.0"E 06 - 08 77 to 15°32'25.8"N, 73°45'04.9"E 32. 6.01.2007 PFG* 15°32'25.8"N, 73°45'04.9"E 06 - 07 85 to 15°30'23.2"N, 73°45'39.6"E 33. 26.01.2007 PFG* 15°30'08.1"N, 73°44'56.5"E 09 - 11 84 to 15°32'58.3"N, 73°44'08.4"E 34. 26.01.2007 PFG* 15°32'58.3"N, 73°44'08.4"E 09 - 12 90 to 15°30'12.7"N, 73°44'29.8"E 35. 26.01.2007 PFG* 15030'13.6"N, 73 044'25.1"E 10 - 12 91 to 15°32'57.5"N, 73 043143.9"E 36. 3.02.2007 PFG* 15029'55.1"N, 73 044'58.7"E 09 -11 95 to 15032'25.2"N, 73 044'22.1"E 37. 3.02.2007 PFG* 15032'25.2"N, 73 044'22.1"E 09 - 11 95 to 15°29'36.0"N, 73°45'02.7"E 38. 19.02.2007 PFG* 15°32'39.3"N, 73 044'17.9"E 09 - 11 96 to 15°30'05.4"N, 73°44'58.1"E 39. 19.02.2007 Aguada bay 15°28'08.8"N, 73°47'09.7"E 04 - 06 96 to 15°27'55.2"N, 73°47'39.6"E 40. 11.03.2007 PFG* 15029'43.2"N, 73°44'39.7"E 11 - 14 145 to 15°33'15.2"N, 73°42'41.6"E 41. 11.03.2007 PFG* 15°33'15.2"N, 73 042'41.6"E 13 -14 120 to 15°30'01.0"N, 73°44'07.7"E 42. 25.03.2007 PFG* 15030'43.1"N, 73°45'40.0"E 06 - 07 60 to 15°32'28.1"N, 73°45'09.4"E 43. 11.04.2007 PFG* 15029'54.7"N, 73°44'44.2"E 11 - 14 100 to 15°32'09.8"N, 73°43'28.4"E 44. 11.04.2007 PFG* 15032'09.8"N, 73 043'28.4"E 12 -14 80 to 15°29'41.7"N, 73°44'16.7"E 45. 25.04.2007 PFG* 15°29'16.7"N, 73°45'50.1"E 08 - 11 105 to 15028'51.9"N, 73 045'45.2"E 46. 25.04.2007 PFG* 15°30'12.8"N, 73°44'55.7"E 08 - 11 75 to 15°32'21.5"N, 73°44'28.3"E 47. 25.04.2007 PFG* 15°29'59.6"N, 73°45'06.1"E 08 - 11 30 to 15029'20.3"N, 73°45'46.0"E 48. 4.05.2007 PFG* 15030'02.6"N, 73°44'54.7"E 10 - 11 90 to 15°30'38.3"N, 73°44'41.7"E 49. 4.05.2007 PFG* 15030'38.3"N, 73°44'41.7"E 10 - 11 130 to 15°30'37.3"N, 73°44'42.7"E 50. 4.05.2007 PFG* 15°30'37.3"N, 73 044'42.7"E 10 -11 110 to 15030'00.1"N, 73°44'51.4"E 51. 16.05.2007 Zuari estuary 15°25'58.1"N, 73°48'09.6"E 03 - 06 62 to 15025'24.9"N, 73°49'53.1"E 52. 29.05.2007 PFG* 15029'40.0"N, 73 045'25.0"E 0.' -11 110 to 15°32'30.9"N, 73°44'08.7"E 53. 29.05.2007 PFG* 15°32'30.9"N, 73°44'08.7"E 09 - 11 105 to 15°30'36.9"N, 73°44'52.3"E 54. 29.05.2007 PFG* 15°30'36.9"N, 73°44'52.3"E 09 - 11 155 to 15°30'45.3"N, 73°44'45.6"E 55. 28.09.2007 Mandovi 15°30'25.8"N, 73°56'57.0"E 02 - 03 60 estuary to 15°30'20.0"N, 73°52'08.3"E 56. 29.09.2007 Zuari estuary 15°26'48.3"N, 73°48'27.4"E 04 - 05 60 to 15°25'59.2"N, 73°50'40.6"E 57. 29.09.2007 Mormugao bay 15°25'16.3"N, 73°51'13.3"E 05 - 06 60 to 15°25'13.8"N, 73°49'39.5"E 58. 3.11.2007 PFG* 15°30'37.8"N, 73°43'06.9"E 15 -17 120 to 15°32'51.2"N, 73°41'56.1"E 59. 3.11.2007 PFG* 15°32'51.2"N, 73°41'56.1"E 15 - 17 80 to 15°30'30.1"N, 73°42'54.5"E 60. 30.11.2007 PFG* 15°30'21.3"N, 73°42'50.5"E 16 -17 120 to 15°33'35.2"N, 73°41'59.4"E 61. 30.11.2007 PFG* 15°33'35.2"N, 73°41'59.4"E 17 -18 120 to 15°30'40.5"N, 73°42'29.3"E 62. 30.11.2007 PFG* 15°30'40.5"N, 73°42'29.3"E 16 - 18 120 to 15°30'45.6"N, 73°42'41.9"E 63. 12.12.2007 Mandovi 15°30'28.0"N, 73°51'35.8"E 04 - 08 7 estuary to 15°30'24.7"N, 73°51'04.9"E 64. 12.12.2007 Aguada bay 15°28'49.5"N, 73°47'23.3"E 05 - 08 40 to 15°28'19.0"N, 73°46'32.3"E 65. 9.01.2008 Mandovi 15°30'25.8"N, 73°56'57.0"E 02 - 03 22 estuary to 15°30'20.0"N, 73°52'08.3"E 66. 9.01.2008 Cumbharjua 15°31'17.1"N, 73°55'57.8"E 03 - 04 16 canal to 15°30'58.6"N, 73°56'22.3"E 67. 9.01.2008 Cumbharjua 15°29'11.7"N, 73°57'19.2"E 04 - 05 20 canal to 15°28'41.1"N, 73°57'03.0"E 68. 9.01.2008 Cumbharjua 15°25'38.0"N, 73°55'38.1"E 03 - 04 06 canal to 15°25'28.6"N, 73°55'36.8"E 69. 9.01.2008 Zuari estuary 15°24'59.7"N, 73°52'38.9"E 03 - 04 34 to 15°25'17.8"N, 73°51'29.1"E 70. 9.01.2008 Zuari estuary 15°25'17.8"N, 73°51'29.1"E 03 - 04 45 to 15°25'22.2"N, 73°51'33.2"E 71. 29.01.2008 PFG* 15°30'12.8"N, 73°43'04.2"E 15 - 17 110 to 15°30'15.0"N, 73°43'03.1"E

72. 29.01.2008 PFG* 15°30'12.0"N, 73°43'04.2"E 1 3 - 17 110 to 15°30'14.7"N, 73°42'09.0"E 73. 29.01.2008 PFG* 15°30'18.1"N, 73°43'04.2"E 16 - 17 85 to 15°30'11.6"N, 73°42'09.8"E 74. 10.02.2008 PFG* 15°31'35.9"N, 73°44'29.9"E 17 - 18 130 to 15°31'36.6"N, 73°44'30.0"E 75. 10.02.2008 PFG* 15°32'00.5"N, 73°44'48.0"E 16 -18 55 to 15°30'53.6"N, 73°44'38.4"E 76. 10.02.2008 PFG* 15°31'00.8"N, 73°44'32.3"E 15 - 17 120 to 15°30'58.9"N, 73°44'38.5"E 77. 23.02.2008 PFG* 15°30'57.8"N, 73°44'39.1"E 15 - 17 140 to 15°32'31.3"N, 73°44'06.8"E 78. 23.02.2008 PFG* 15°32'25.4"N, 73°43'55.2"E 15 - 17 165 to 15°30'32.8"N, 73°43'57.0"E 79. 08.03.2008 PFG* 15°30'51.2"N, 73°42'13.6"E 18 - 20 160 to 15°30'20.1"N, 73°41'14.0"E 80. 08.03.2008 PFG* 15°30'20.1"N, 73°41'14.0"E 18 - 20 170 to 15°30'54.0"N, 73°42'21.0"E 81. 19.03.2008 PFG* 15°30'08.7"N, 73°43'00.9"E 16 - 17 165 to 15°33'39.2"N, 73°41'46.4"E 82. 19.03.2008 PFG* 15°33'17.8"N, 73°41'46.5"E 16- 17 130 to 15°30'17.5"N, 73°42'52.8"E 83. 23.04.2008 PFG* 15°30'54.7"N, 73°45'31.9"E 06 - 09 110 to 15°31'45.3"N, 73°44'38.9"E 84. 23.04.2008 PFG* 15°31'41.5"N, 73°44'54.8"E 06 - 09 90 to 15°31' 17.4"N, 73°45'00.0"E 85. 23.04.2008 PFG* 15°31'29.0"N, 73°44'54.2"E 08 - 09 102 to 15°31'10.8"N, 73°45'01.2"E 86. 30.04.2008 PFG* 15°31'00.0"N, 73°44'55.8"E 09 - 11 110 to 15°31'09.7"N, 73°44'34.7"E 87. 30.04.2008 PFG* 15°32'00.0"N, 73°45'05.1"E 06 - 08 85 to 15°31'56.0"N, 73°45'12.0"E 88. 30.04.2008 PFG* 15°31'56.0"N, 73°45'12.0"E 07 - 11 102 to 15°29'24.3"N, 73°45'43.4"E 89. 10.05.2008 PFG* 15°30'51.9"N, 73°44'55.6"E 09 — 11 136 to 15°31'00.5"N, 73°44'42.7"E 90. 10.05.2008 PFG* 15°30'56.5"N, 73°44'41.0"E 10 — 11 120 to 15°30'54.6"N, 73°44'43.4"E 91. 10.05.2008 PFG* 15°30'54.6"N, 73°44'43.4"E 09 — 11 136 to 15°31'12.2"N, 73°44'59.7"E 92. 28.11.2008 PFG* 15°31'15.0"N, 73°45'23.2"E 04 — 05 80 to 15°32'13.5"N, 73°45'13.2"E 93. 28.11.2008 PFG* 15°31'32.4"N, 73°45'07.0"E 04 — 05 35 to 15°32'13.5"N, 73°45'12.1"E 94. 28.11.2008 PFG* 15°32'28.3"N, 73°44'21.1"E 04 — 06 55 to 15°30'34.1"N, 73°44'43.5"E 95. 28.11.2008 PFG* 15°30'35.9"N, 73°45'38.0"E 04 — 06 105 to 15°30'44.3"N, 73°45'34.7"E

*PFG — Potential Fishing Grounds off Goa between Aguada bay and Baga estuary bay, five in Zuari estuary, three each in Mandovi estuary and Cumbharjua canal and two in Aguada bay) in view of restricted fishing activity due to prohibition on fishing in inland waters. Geographical position of each sampling station was recorded with a

12 — Channel Geographical Positioning System (GPS) and sampling depth for the same was obtained from the Naval Hydrographic Chart No. 2022. Trawl nets with mesh sizes of 15 mm (mouth end) and 9 mm (cod end) were towed approximately at a speed of 2 knots (4 km.11 -1). The sampling duration was for one to three hours in the nearshore region, whereas, in the estuaries the irregular bottom topography restricted the trawl sampling to a maximum of one hour. In addition, samples from the estuarine embayment those inaccessible to bottom trawls were obtained by operating beach seine (one hour operation) along Betim (15°30'18"N, 73°49'52"E) during May, 2005 and three in the vicinity of the Mormugao Port Trust (15°24'16"N, 73°48'56"E) during December, 2005 and September, 2006 (Table 2.2). Fish samples were also collected from local fishermen operating gill nets near the mouth of the Mandovi estuary in December, 2005 to assess the occurrence of these species in the estuarine embayment. Estuarine crabs those available in meagre quantity in the trawl catches were obtained from the estuarine embayment using crab traps and also obtained from commercial outlets.

The trawl catch obtained was first examined for species composition and the same was recorded. Subsequently, five sub — samples were randomly picked from the catch. Out of the ninety five trawl samplings carried out, adequate catch in excess of

30 kg was obtained only from sixty eight trawls, hence sub — sampling was done only for these hauls. The specimens those were found to be uncommon or rare were picked out separately, put on ice prior to transportation to the laboratory for detailed examination. Similarly, biological samples obtained from beach seines, crab traps and

10 Table 2.2. Details of shore sampling (beach seine and crab trap) carried out along the bay — estuarine waters of Goa

Sr. Sampling Sampling area Geographical Position Sampling Sampling No. date depth (m) duration (mm) 1. 8.05.2005 Betitn bay° 15°30'18.0"N, 73°49'52.0"E 0: — 02 60 2. 8.05.2005 RND jetty 15°30'21.0"N, 73°49'49.0"E 01 — 02 60 3. 13.12.2005 Vasco bay# 15°24'47.0"N, 73°48'38.5"E 01 — 02 60 4. 13.12.2005 Betim* 15°30'18.0"N, 73°49'52.0"E 01 —02 60 5. 13.12.2005 Vasco — 15°24'30.0"N, 73°50'06.0"E 01 — 02 60 Kharivadem# 6. 26.09.2006 Vasco — 15°24'30.0"N, 73°50'06.0"E 01 - 02 60 Kharivadem# • teach seine operation *Crab trap gill net collection were also examined for species composition, stored in ice and sent to the laboratory for detailed identification.

2.3. Taxonomic identification and morphometry

At the laboratory, the samples were photographed using a 7.2 mega pixel digital camera (SONY DSC 5750, 3X optical zoom). In addition, minute details were recorded by Camera lucida diagrams using an Olympus SZX — DA 3M01330 microscope.

Subsequently, the samples were identified using conventional taxonomic methods involving phenotypic analyses such as morphology, colour, texture patterns, meristic counts (Table 2.3) and morphological measurements (Table 2.4) up to the nearest 0.01 cm using vernier callipers. The above identification was aided by published taxonomic literature for the respective faunal groups: fin fish (Day, 1876 —

1878; Lindberg, 1973; Fischer and Whitehead, 1974; Fischer and Bianchi, 1984;

Talwar and Kacker, 1984; Talwar and Jhingran, 1991); prawns (George, 1980; Kurian and Sebastian, 1986; Chan, 1998); stomatopods (Manning, 1998); brachyuran crabs

(Alcock 1895, 1896, 1899a, 1900; Leene, 1938; Chhapgar, 19f,7; Sakai, 1976;

Sethuramalingam and Khan, 1991; Wee and Ng, 1995; Jeyabaskaran et al., 2002); anomuran crabs (Khan, 1992); molluscs (Silas et al., 1983; Roper et al., 1984;

Mookherjee, 1985; Rao and Rao, 1993; Wilson, 1994; Apte, 1998; Rajagopal et al.,

1998); echinoderms (Clark and Rowe, 1971); sea snakes (Rasmussen, 2001). In addition, internet websites such as Fishbase (Froese and Pauly, 2010), Sealifebase

(Palomares and Pauly, 2010), and Hardy's internet guide to Marine Gastropods

(Hardy, 2010) were referred for species identification.

11 Table 2.3. List of meristic counts used in the identification of demersal marine fauna

Sr. No . roup Meristic counts (parameter) 1. Elasmobranchs No. of spines on tail No. of papillae on floor of buccal cavity 2. Teleosts No. of dorsal fin spines No. of dorsal fin rays No. of anal fin spines No. of anal fin rays No. of pectoral fin rays No. of pelvic fin spines No. of pelvic fin rays No. of caudal fin rays (Cynoglossidae) No. of gill rakers on the first gill arch No. of nostrils No. of pores on chin (Sciaenidae, Haemulidae) No. of barbels on chin (Ariidae, Mullidae) No. of scutes on belly (Clupeidae, Engraulidae) No. of lateral lines (more than one in Cynoglossidae) No. of scales on lateral line No. of scale rows between dorsal fm origin and lateral line No. of branchiostegal rays 3. Stomatopods No. of spines on carina of abdominal segment _ No. of teeth on dactylus of raptorial claw 4. Prawns No. of spines on rostrum 5. Brachyuran crabs No. of teeth on antero-lateral margin of carapace No. of spines on merus of cheliped No. of spines on carpus of cheliped No. of spines on manus of cheliped 6. Anomuran crabs No. of pleopod pairs on abdomen 7. Cephalopods - 8. Gastropods No. of spire whorls No. of teeth on outer lip of aperture No. of denticles on inner lip of aperture No. of varices No. of spines on anterior siphonal canal No. of spiral cords, tubercles etc. 9. Bivalves 10. Echinoderms No. of ambulacral plates (sea urchin) No. of buccal plates on peristome (sea urchin) 11. Sea snakes (Reptiles) - Table 2.4. List of moThometric parameters used in the identification of demersal marine fauna

Sr. No. Faunal group Morphometric parameter 1. Elasmobranchs Disc width . Disc length Snout / pre-oral length Inter — orbital width Inter — spiracular width Tail length 2. Teleosts Total length Standard length Fork length Body depth Head length Orbital diameter Pre — orbital head length Inter — orbital width Snout length 3. Stomatopods Total length 4. Prawns Carapace length Rostrum length Width of adrostral carina Length of adrostral groove 5. Brachyuran crabs Carapace width Carapace length Frontal width Inter — orbital space Length of chelipeds Length of pereiopods 6. Anomuran crabs Total length Carapace length Carapace width Rostral length 7. Cephalopods Dorsal mantle length (cuttle fish, squids) Fin length (squids) Fin width (squids) Ann length (cuttle fish) 8. Gastropods Total shell length Spire height Body whorl length 9. Bivalves Shell height 10. Echinoderms Horizontal head diameter (sea urchin) Vertical test diameter (sea , urchin) Peristome diameter (sea urchin) Diameter of apical system (sea urchin) Periproct diameter (sea urchin) Arm length (sea star) A 11. Sea snakes (Reptiles) Snout vent length Total length Morphometric analysis involved measurement of morphological parameters of the biological specimens and subsequent comparison with published data (Froese and

Pauly, 2010; Palomares and Pauly, 2010).

2.4. Sample preservation

Representative specimens of fin fish, molluscs and other faunal groups were preserved using 5 % formalin, whereas crustaceans were preserved in 5 % buffered formalin (buffered with hexamethylene tetramine to prevent fragmentation of appendages). These are stored in pre — labelled transparent plastic bottles and deposited as reference vouchers at the Marine Biology laboratory, Department of

Marine Sciences, Goa University.

2.5. 18S rDNA sequencing

18S rDNA sequencing was employed to ascertain species level identity of one form of the mud crab, Scylla and the puffer fish, Tetraodon. The protocol for the above technique comprised excision of about 2 g of tissue from the specimen, followed by preservation in absolute alcohol. The preserved samples were sent to

Chromous Biotech Ltd., a biotechnology — related company at Bengaluru, India for

18S rDNA extraction. The total genomic DNA was extracted using Chromous Animal

Tissue genomic DNA isolation kit RKTO9 following the manufacturer's protocol and genome was amplified using Polymerase Chain Reaction (PCR) technique using two primers (Forward primer: 5'-TICTCCACCAACCACAA(A/G)GA(C/T)AT(C/T)GG-

3' and Reverse primer: 5'-CACCTCAGGGTGTCCGAA(A/G)AA(C/T)CA(A/G)AA-

3'). The PCR reaction involved initial denaturation at 94°C for five minutes, followed by a reaction cycle (94°C for thirty seconds, 55°C for thirty seconds, 72°C for one

12 minute) repeated thirty five times with a final extension step of 72°C for fifteen minutes. The PCR product was gel purified using Chromous Gel Extraction Kit

(RKT33) and sequenced. Subsequently, the Basic Local Alignment Search Tool

(BLAST) of the NCBI (Gene Bank) website was employed to obtain the sequence alignment and phylogenetic position.

2.6. Data compilation and processing

2.6.1. Faunal composition

All taxa observed during the present study were divide(' into four broad taxonomic groups namely fin fish, crustaceans, molluscs and other fauna.

Subsequently, a list of the above taxa was tabulated along with the status of reporting from the study area.

2.6.2. Faunal abundance (numbers) and weight

Data on abundance (numbers) and weight of constituent species from five sub

— samples of each trawl haul were computed as follows.

= [x, + x2 + x3 + x4 + xs] / 5 where 'x' denotes abundance (numbers) or weight of a taxon in a sub — sample.

The above data was standardized to sixty minute — trawl in view of the variability in trawling duration, by converting values for variable durations to per sixty minute — haul using the following equation.

X' = ER x 60] / tact where, toe) — actual trawling duration (in minutes)

X — data value (abundance or weight) for tact

X' — data value for sixty minute haul

13 Thereafter, the data was segregated into four principal faunal groups namely fin fish, crustaceans, molluscs and other fauna, as well as their sub . — groups to determine their contribution to the total trawl catch. In addition, the monthly trends in abundance (numbers) and weight of the principal faunal groups and their sub — groups were computed. Comparisons of abundance (numbers) and weight of faunal groups by month and season were analyzed by two — way Analysis of Variance (ANOVA; Sokal and Rohlf, 1987), and compared with F — table (P = 0.05 and 0.001 levels of significance).

2.6.3. Species diversity indices

The present study employed the following four indices to determine the diversity among the demersal marine fish along Goa coast.

2.6.3a. Shannon — Wiener's diversity index

This statistical measure was proposed by Shannon and Wiener (1963) to measure not only the number of different species but also to study their relative proportion and distribution in a sample.

S H = pi x ln (pi) i =1 where, S — total number of species

pi — the relative abundance of species "i" in a sub — sample, calculated as pi= (ni / N) where, ni — number of individuals of species "i"

N — total number of individuals of all species

14 2.6.3b. Heip's evenness index

This statistical measure was proposed by Heip (1974) to measure the relative abundance of individuals of a particular species in a sample.

E = [H'-1] / [S-1] where, H' — Shannon — Wiener diversity function

S — total number of species

2.6.3c. Margalef's species richness

This statistical measure was proposed by Margalef (1968) to measure species richness.

SR = [S — 1] / ln (N) where, S — number of species

N — total number of individuals

2.6.3d. Simpson's dominance index

This statistical measure was proposed by Simpson (1949) and measures biodiversity based on the probability that two individuals randomly selected from a sample will belong to the same species (or other higher taxon).

D=E pi2 where, p1— the relative abundance of species "i" in a sub — sample, calculated as pi= (n1 / N) where, n1 — number of individuals of species "i"

N — total number of individuals of all species

15 2.6.4. Cluster analysis

Cluster analysis is a statistical technique used to assign objects or data into groups (clusters) those exhibit natural groupings and differ from other groups. The present study employed two techniques namely Dendrogram plotting and non — metric Multi — Dimensional Scaling (nMDS) to assess the faunal associations within the demersal fish community.

In view of this, data was normalized using the fourth — rcot transformation function to reduce the influence of aberrant high values and weigh the contributions of both common and rare taxa (Ansari et al., 2003). The above exercise was carried out at two levels namely "families" and "species". Altogether, sixty three "families" were selected for cluster analysis. However, in view of the voluminous data in case of the "species" level, only the most frequently occurring species (N = 38) representing the most common faunal groups were selected. Thereafter, the above data were converted into a lower triangular matrix using the Bray — Curtis similarity coefficient

(Bray and Curtis, 1957) and subjected to Dendrogram plotting ane nMDS using the

Plymouth Routines In Multivariate Ecological Research (PRIMER) version 5 computer program (Clarke and Gorley, 2001).

2.6.5. Functional guilds

The "functional guilds" concept (Elliot et al., 2007) employs ecological parameters such as estuarine use, vertical distribution, substratum preference and feeding habits (Table 2.5) to fish community structure. The ecological requirements of the demersal species such as habitat, migration type and diet preference (Froese and Pauly, 2010; Palomares and Pauly, 2010) were obtained to explain the faunal clusters following the criteria given by Elliot et al. (2007).

16 Table 2.5. Functional guilds used to categorize the demersal marine fauna along Goa coast

Sr. Functional Ecological category Definition No. guild 1. Estuarine Estuarine resident Species that completes its entire life cycle in the estuary use (ER) Estuarine migrant Species with estuarine adults; larval phase outside the (EM) estuary Marine migrant (MM) Species that spawns at sea, juveniles enter estuaries in large numbers Marine straggler (MR) Species that spawns at sea, occurs near the estuary mouth at 35 PSU salinity and enters estuaries in very low numbers Anadromous (AN) Species that undergoes its greatest development at sea and migrates to freshwater environment for spawning Semi — anadromous Species that undergoes its development at sea, and (SA) migrates up to the upper estuary for spawning Catadromous (CA) Species that undergoes its development in freshwater environment and migrates to the sea for spawning Semi — catadromous Species that undergoes its development in freshwater (SC) environment and migrates only to brackish water for spawning Amphidromous (AM) Species that migrates between the freshwater and marine environments; migration not related to reproduction Oceanodromous (OC) Species that migrates exclusively within the marine environment for reproduction 2. Vertical Pelagic Species that lives in the main water column distribution Benthic Species that lives on or within tli... substratum Demersal Species that lives at the sediment — water interface 3. Substratum Soft bottom (F) Species that inhabits fine sediment (mud / silt) preference sub stratum Sand (S) Species that inhabits sandy substratum . Rough (R) Species that inhabits rough (rocky) substratum Mixed (M) Species that inhabits soft / sandy substratum interspersed with submerged rocks or coral rubble Vegetation (V) Species that lives among submerged vegetation 4. Feeding Phytoplanktivore (PP) Species that feeds on phytoplankton habits Zooplanktivore (ZP) Species that feeds on zooplankton Herbivore (HV) Species that feeds on submerged vegetation Omnivore (OV) Species that feeds on macrophyte, epifauria and infauna Piscivore (PV) Species that feeds on nektonic vertebrates (fishes) Zoo — benthivore (ZB) Species that feeds on invertebrates associated with the substratum Opportunistic / Species that feeds on a wide variety of prey and cannot Miscellaneous be included in any of the above categories

4C 2.6.6. Determination of catch rate

Computation of catch rate (kg.hr - 1) of the nearshore waters up to 25 m depth was done by first converting the weight of individual species per sample (expressed in kg) to the total trawl weight using the following equation.

Y = [Xi x Xt] / Xs where, Xs — weight of sample "s"

— weight of individual species "i" in sample "s"

Xt — weight of entire trawl haul "t"

Thereafter, the weight of individual species per trawl (Y) was converted to sixty minute — haul to yield the catch rate (Y', expressed in kg.hr -1) using the following equation.

Y' = [Y x 60] / tact where, tact — actual trawling duration (in minutes)

Y — data value (weight) for tact

Y' — data value (weight) for sixty minute — haul

The above data was categorized into fifteen fish groups namely .stomatopods, prawns, elasmobranchs, clupeoids, sciaenids, lactariids, catfishes, leiognathids, ribbon fishes, flatfishes, carangids, pomfrets, perches, cephalopods and miscellaneous following Prabhu and Dhawan (1974) to facilitate comparison with the latter.

17 Chapter 3. Demersal faunal community structure off Goa 3.1 Introduction

The community structure concept focuses the interactions among species and their environments including trophic networks (Pianka, 1973; Cody, 1974) and enables better understanding not only of the species, but also the associated biological processes those regulate the community. These studies comprise (a) qualitative analyses involving species composition and their ecological categorization, and (b) quantitative analyses pertaining to faunal abundance, weight, diversity and spatio — temporal variations among these parameters. The above information is further employed to resolve issues related to species interactions and associations. Moreover, the ever — growing impact of anthropogenic activities on the coastal waters enforces continuous assessment to take a holistic view of such ecosystem.

In view of this, a comprehensive review of literature was carried out to explore various aspects of the demersal marine community structure in coastal ecosystems across various bioregions of the world.

3.2. Literature review

Published literature pertaining to the demersal marine fish communities indicate that extensive studies have been carried out in the major bay — estuarine and coastal ecosystems (Elliot et al., 2007). A majority of these studies have been carried out in the Indo — Western Pacific regions (Rainer and Munro, 1982; Wallace et al.,

1984; Blaber et al., 1985, 1989, 1994, 1995; Chan and Liew, 1986; McManus, 1986,

1989, 1996; Little et al., 1988; Qui, 1988; Whitfield, 1988; Wright, 1988; Kihara and

Itosu, 1989; Loneragan et al., 1989; Blaber and Milton, 1990; Chong et al., 1990;

Potter et al., 1990; Robertson and Duke, 1990; Watson et al., 1990; Chittima and

Wannakiat, 1992; Federizon, 1992; Ansari et al., 1995, 2003; Harrison and Whitfield,

18 1995, 2006; Martin et al., 1995; Bianchi, 1996; Laroche et al., 1997; Adjeroud et al.,

1998; Potter and Hyndes, 1999; Kuo et al., 2001; Alias, 2003; Campos, 2003;

Hajisamae et al., 2003; Harrison, 2003; Khongchai et al., 2003; Mustafa, 2003;

Nurhakim, 2003; Srinath et al., 2003; Lugendo et al., 2007; Chen et al., 2009;

Haicheng and Weiwei, 2009; Hajisamae, 2009; Yemane et al., 2010). The above studies were focused at describing the community structure of demersal fishes of both pristine and highly impacted coastal habitats through spatio — temporal and trophic analyses of the constituent fish populations. In addition, a few of these studies

(Loneragan et al., 1989; Martin et al., 1995; Kuo et al., 2001; Ansari et al., 2003;

Lugendo et al., 2007; Haicheng and Weiwei, 2009) attempted to elucidate the role of environmental factors on the community structure per se and suggested that environmental anomalies play vital role in the composition of biotic communities.

Studies from Europe and the Mediterranean region (Chesney and Iglesias,

1979; Iglesias, 1981; Claridge et al., 1986; Pomfret et al., 1991; Potter et al., 1986,

1997; Thiel et al., 1995; Elliot and Dewailly, 1995; Marshall and Elliot, 1998; Ratz,

1999; Koutrakis et al., 2000; Mathieson et al., 2000; Gordo and Cabral, 2001; Lobry et al., 2003; Prista et al., 2003; Labropoulou and Papaconstantinou, 2005; Akin et al.,

2005; Leitao et al., 2007; Selleslagh and Amara, 2008) dealt with the demersal faunal assemblages of temperate estuaries and boreal shelf waters and revealed that salinity, temperature and depth played a major role in structuring species assemblages of these environments.

Albaret et al. (2004), Guillard et al. (2004) and Simier et al. (2006) intensively studied the spatio — temporal variability in fish diversity and distribution within the

Gambia estuary (Atlantic coast of Africa) in relation to environmental variables and

19 suggested the use of this estuarine system as a reference to study the effects of climatic perturbations on estuarine fish communities.

Published literature from the Atlantic coast of the Americas (Hoff and Ibara,

1977; Yafiez — Arancibia et al., 1980; Sedberry and van Dolah, 1984; Chao et aL,

1985; Stoner, 1986; Paiva — Filho et al., 1987; Villarroel, 1994; Aratijo et al., 1998,

2002; Chaves and Correa, 1998; Garcia et al., 1998; Arailjo and Costa de Azevedo,

2001; Cousseau et al., 2001; Nordlie, 2003; Nagelkerken and van der Velde, 2004;

Barletta et al., 2005, 2008; Gonzalez — Castro et al., 2009) revealed that salinity was the dominant factor determining species composition, abundance and diversity of demersal fish communities. Studies on the community structure of demersal fish from the Pacific coast of the Americas (Allen and Horn, 1975; Allen, 1982; Bartels et al.,

1983; Amezcua — Linares et al., 1987; Yoklavich et al., 1991; Monaco et al., 1992;

Rojas et al., 1994; Wolff, 1996; Gonzalez — Acosta et al., 2005) suggested that physico — chemical (salinity, depth, temperature and proximity to the sea) and biological factors (food availability, reproduction and migration patterns) collectively influenced the community structure.

Recent community structure studies from estuaries and coastal waters reveal the use of the "ecological guild" concept. This identifies subsets within species assemblages those have high potential for competition (Simberloff and Dayan, 1991) along with identification and prediction of species responses to environmental variation given the known set of environmental parameters for that species (Austen et al., 1994). This concept was successfully used to analyze estuarine fish communities of some of the major estuaries and nearshore coastal waters of the Indo — Pacific

(Potter et al., 1990; Blaber, 1991; Harrison and Whitfield, 1995; Potter and Hyndes,

1999; Hajisamae et al., 2003; Haicheng and Weiwei, 2009; Hajisamae, 2009) and

20 Atlantic (Haedrich, 1983; Potter et al., 1986; Elliot and Dewailly, 1995; Mathieson et al., 2000; Cousseau et al., 2001; Lobry et al., 2003; Nordlie, 2003; Albaret et al.,

2004; Nagelkerken and van der Velde, 2004; Leitdo- et al., 2007; Gonzalez — Castro et al., 2009) regions. Elliot et al. (2007) in an attempt to apply the guild concept to adjacent marine and freshwater niches, collectively known as "transitional waters", refined previously used terminologies and their definitions in order to facilitate unification of these terms for use across geographical boundaries.

It is imperative that these studies focused only at demersal ichthyofauna, thus creating lacunae in understanding the role of invertebrate components. In contrast, only few studies (De Ben et al., 1990; Maes et al., 1998; Ungaro et al., 1999; Akin et al., 2003; Chesoh et al., 2009) attempted to describe both demersal fin fish and invertebrate communities. De Ben et al. (1990) reported spatio — temporal variations in fin fish and crustacean distribution and abundance from the Yaquina Bay, Oregon

(USA) and attributed the same to variations in salinity and temperature. Macs et al.

(1998) reported marked seasonal variations in the fin fish and crustacean community of the Zeeschelde estuary, Belgium and suggested that variation in predation risk played an important role in determining such variations. Ungaro et al. (1999) attempted to describe the fin fish, crustacean and cephalopod assemblages along the

South Adriatic coast of the Mediterranean Sea and revealed a strong association between fin fish distribution and depth, whereas the influence of depth on invertebrates was comparatively less. Akin et al. (2003) reported marked seasonal and spatial variations in fin fish and crustacean assemblages from the Mad Island Marsh estuary, Texas (USA), and attributed the same to the combined effects of faunal migration patterns and species — specific response to depth and local environmental variation. Chesoh et al. (2009) attempted to study the demersal fin fish, crustacean

21 and cephalopod communities of the coastal Songkhla lake, Thailand, and formulated a model pertaining to the faunal assemblages of the region based on variations in weights of catches along the lake with applications in fishery landings. The present study makes an attempt to inventorize the demersal fauna along the estuarine — coastal system. Further, efforts are being made to examine the data to compute various parameters and the same has been subjected to statistical analysis to provide better insight into the species composition and their possible interactions within the system.

3.3. Results

3.3.1 Species composition

The present study based on the data collected from the bay — estuarine and coastal potential fishing grounds of Goa, central west coast of India up to 25 m depth attempts to elucidate the demersal marine faunal diversity of the region, inclusive of common and rare taxa. The above exercise revealed altogether two hundred and four taxa belonging to five phyla, ten classes, two clades, twenty seven orders, ninety nine families and one hundred and fifty nine genera. These taxa were further categorized into four major faunal groups (Fig. 3.1). Among these, the fin fish (139 taxa) were represented by the most species, followed by crustaceans (35), and molluscs (24); minor faunal groups namely sea snakes (03), echinoderms (02) and cnidarians (01) were clubbed together as 'other fauna'. Among these, two hundred and one taxa were identified up to the species level (Table 3.1).

Preliminary analysis of the demersal faunal groups observed during the present study revealed that crustaceans, molluscs, echinoderms, sea snakes and

resident fin fish were represented by juveniles as well as adult individuals (Table 3.2).

On the other hand, juveniles and young recruits dominated the migrant fin fish

22 Other fauna* Molluscs 6 24

Crustaceans 35

Fin fish 139

*Echinoderms (02), Cnidaria (01), Sea snakes (03)

Fig. 3.1. Taxa composition of demersal faunal groups observed during the present study Table 3.1. List of demersal fauna identified during the present study from Goa, central west coast of India

Sr. Taxa Reports from No.. Goa coast

Fin fish 1. Pastinachus sephen (ForskAl, 1775) G16 2. Neotiygon kuhlii (Muller & Henle, 1841) Present study 3. Himantura gerrardi (Gmelin, 1789) Present study 4. Himantura walga (Miller & Henle, 1841) Present study 5. Aetobatus flagellum (Bloch & Schneider, 1801) Present study 6. Rhinobatos obtusus Muller & Henle, 1841 Present study 7. Glaucostegus granulatus (Cuvier, 1829) G16, G23 8. Chiloscyllium griseum (Muller & Henle, 1838) G2 9. Scoliodon laticaudus (Muller & Henle, 1838) G2, G4, G14, G16, G17, G23 10. Sardinella longiceps (Valenciennes,1847) G2, G16, G19, G23 11. Sardinella brachysoma Bleeker, 1852 ?? 12. Escualosa thoracata (Valenciennes, 1847) G2, G16, G19 13. Nematalosa nasus (Bloch, 1795) G2, G3, G16 14. Dussunzieria acuta (Valenciennes, 1847) G2, G14, G16, G17, G23 15. Opisthopterus tardoore (Cuvier, 1829) G2, G4, G14, G16, G17, G19 16. Chirocentrus dorab (Forskal, 1775) G2, G14, G16, G17, G19, G23 17. Pellona ditchela Valenciennes, 1847 G2, G4, G14, G17 18. Coilia dussumieri (Valenciennes, 1848) G14, G16, G17 19. Thryssa malabarica (Bloch, 1795) G2, G3 20. Thryssa mystax (Bloch & Schneider, 1801) Present study 21. Thryssa puraya (Hamilton, 1822) G2, G16 22. Thryssa setirostris (Broussonet, 1782) Present study 23. Thryssa dussumieri (Valenciennes, 1848) G2, G16 24. Stolephorus commersonnii (Lacepede, 1803) G16, G22 25. Stolephorus baganensis Hardenberg, 1933 G2 26. Hyporhamphus quoyi (Valenciennes, 1847) G2 27. Hippocan2pus kuda Bleeker, 1852 Present study 28. Mugil cephalus Linnaeus, 1758 G19 29. Siganus canaliculatus (Park, 1797) G2, G16 30. Scatophagus argus (Linnaeus, 1766) G2, G3, G16, G23 31. Ambassis gymnocephalus (Lacepede, 1802) G2, G3, G16 32. Apogon fasciatus (White, 1870) Present study 33. Archamia bleekeri (Gunther, 1859) Present study 34. Alectis indicus (Riippell, 1830) G2, G23 35. Alepes djedaba (Forskal, 1775) G2, G23 36. Atropus atropos (Bloch & Schneider, 1801) G2, G14, G17, G23 37. Carangoides malabaricus (Bloch & Schneider, 1801) G2, G14, G17, G23 38. Caranx ignobilis (Forskal, 1775) G23 39. Decapterus russelli (klippen, 1830) G23 40. Megalaspis cordyla (Linnaeus, 1758) G2, G14, G16, G17, G23 41. Parastromateus niger(Bloch, 1795) G2, G4, G16, G23 42. Scomberoides commersonnianus (Lacepede, 1801) Present study 43. Sconzberoides lysan (Forskal, 1775) G2, G23

44. Rachycentron canadum (Linnaeus, 1766) G2 9 G16 7 023 45. Heniochus acuminatus (Linnaeus, 1758) Present study 46. Drepane punctata (Linnaeus, 1758) G2, G16, G23 47. Drepane longimana (Linnaeus, 1758) Present study 48. Echeneis naucrates (Linnaeus, 1758) G23 49. Gerres eiythrourus (Bloch, 1791) Present study 50. Gerres filamentostes (Cuvier, 1829) G2, G16 51. Gerres limbatus Cuvier, 1830 G16 52. Gerres longirostris Lacepede, 1801) Present study 53. Lactarius lactarius (Bloch & Schneider, 1801) 01, G2, G4, G14, G16, G17, G19, G23 54. Gazza minuta (Bloch, 1795) Present study 55. Photopectoralis bindus (Valenciennes, 1835) G4, G14, G17, G23 56. Nuchequula blochii (Valenciennes, 1835) G2, G16 57. Leiognathus brevirostris (Valenciennes, 1835) Present study 58. Leiognathus daura (Cuvier, 1829) G2, G16 59. Eubleekeria splendens (Cuvier, 1829) G2, G4, G14, G17 60. Secutor insidiator (Bloch, 1787) G2, G4, G16, G23 61. Lutjanus johnii (Bloch, 1792) G2, G3, G14, G16, G17 62. Lutjanus argentimaculatus (Forskal, 1775) G2, G14, G16, G17, G19 63. Caesio tuning (Bloch, 1791) Present study 64. Mene maculata (Bloch & Schneider, 1801) G16, G23 65. Monodactylus argenteus (Linnaeus, 1758) Present study 66. Upeneus vittatus (Forskal, 1775) G2, G16 67. Nemipterus japonicus (Bloch, 1791) G2, G16, G19, G23 68. Parascolopsis townsendi Boulenger, 1901 Present study 69. Pempheris molucca Cuvier, 1829 Present study 70. Eleutheronema tetradactylum (Shaw, 1804) G16, G19, G23 71. Filimanus heptadactyla (Cuvier, 1829) G2, G16, G23 72. Daysciaena albida (Cuvier, 1830) G2, G14,G17 73. Dendrophysa russelii (Cuvier, 1829) Present study 74. Johnius spp. G2, 014,017, G19, G23 75. Otolithes ruber (Bloch & Schneider, 1801) G2, G4, G14, G17, G16, G23 76. Pennahia anea (Bloch, 1793) G2, G14, G17 77. Epinephelus coioides (Hamilton, 1822) Present study 78. Epinephelus diacanthus (Valenciennes, 1828) G2, G16, G23 79. Epinephelus erythrurus (Valenciennes, 1828) Present study 80. Sillago sihanza (ForskAl, 1775) G2, G14, G16, G17, G19 81. Acanthopagrus berda (ForskAl, 1775) G2, G16 82. Sparidentex hasta (Valenciennes, 1830) Present study 83. Pomadasys hasta (Bloch, 1790) Gl, G2, G3, G16 84. Pomadasys fitrcatus (Bloch & Schneider, 1801) Present study 85. Pomadasys maculatus (Bloch, 1793) G2, G16, G23 86. Plectorhinchus gibbosus (Lacepede, 1802) Present study 87. Plectorhinchus schotaf(Forskal, 1775) Present study 88. Terapon jarbua (Forskal, 1775) G2, G16, G23 89. Terapon theraps Cuvier, 1829 G2, G16 90. Terapon puta Cuvier, 1829 G2, G16 91. Scomberomorus guttatus (Bloch & Schneider, 1801) G2, G16, G23 92. Rastrelliger kanagurta (Cuvier, 1816) G2, G16, G19 G23 93. Trichiurus lepturus Linnaeus, 1758 G2, G14, G17, G23 94. Sphyraena putnamae Jordan & Seale, 1905 G23 95. Sphyraena obtusata Cuvier, 1829 G16, G23 96. Pampus argenteus (Euphrasen, 1788) G2, G4, G14, G16, G17, G19, G23 97. Pampus chinensis (Euphrasen, 1788) G2, G4, G14, G16, G17, G19, G23 98. Glossogobis giuris (Hamilton, 1822) G2, G3, G6, G16 99. Paratrypauchen microcephalus (Bleeker, 1860) G2 100. Trypauchen vagina (Bloch & Schneider, 1801) G2, G16 101. Odontamblyopus rubicundus (Hamilton, 1822) G2 102. Yongeichthys criniger (Valenciennes, 1837) Present study 103. Parachaeturichthys polynema (Bleeker, 1853) Present study 104. Oxyurichthys paulae Pezold, 1998 Present study 105. Callionymus japonicus Houttuyn, 1782 Present study 106. Callionymus sagitta Pallas, 1770 Present study 107. Eurycephalus carbunculus (Valenciennes, 1833) Present study 108. Grammoplites scaber (Linnaeus, 1758) G2, G16 109. Trachicephalus uranoscopus (Bloch & Schneider, 1801) G9, G16 110. Cynoglossus arel (Bloch & Schneider, 1801) G14, G23 111. Cynoglossus dubius Day, 1873 G2, G14, G17 112. Cynoglossus puncticeps (Richardson, 1846) G2 113. Cynoglossus macrostomus Norman, 1928 G2, G14, G17 114. Paraplagusia bilineata(Bloch, 1785) G2 115. Synaptura albomaculata Kaup, 1858 Present study 116. Synaptura commersonnii (Lacepede, 1802) G4, G16 117. Solea ovata Richardson, 1846 G2, G16 118. Pseudorhombus triocellatus (Bloch & Schneider, 1801) G2, G16 119. Pseudorhombus arsius (Hamilton, 1822) G2, G14, G17 120. Triacanthus biaculeatus (Bloch, 1786) Present study 121. Lactoria cornuta (Linnaeus, 1758) G16 122. Lagocephalus spadiceus (Richardson, 1845) G2 123. Takifugu oblongus (Bloch, 1786) Present study 124. Arothron immaculatus (Bloch & Schneider, 1801) Present study 125. Chelonodon patoca (Hamilton, 1822) G2 126. Tetraodon fluviatilis fluviatilis Hamilton, 1822 Present study 127. Colletteichthyes dussumieri (Valenciennes, 1837) G2 128. Bregmaceros mcclellandi Thompson, 1840 G16 129. Arius maculatus (Thunberg, 1792) G2, G14, G17 130. Arius subros tratus Valenciennes, 1840 Present study 131. Nemapteryx caelata (Valenciennes, 1840) Present study 132. Netuma thalassina (Ruppell, 1837) G2, G14, G17 133. Mystus gulio (Hamilton, 1822) G2 134. Plotosus lineatus (Thunberg, 1787) G14, G16, G17 135. Pisodonophis cancrivorus (Richardson, 1848) G2 136. Muraenesox bagio (Hamilton, 1822) Present study 137. Muraenesox cinereus (Forskal, 1775) G2, G23 138. Saurida tumbil (Bloch, 1795) G2, G14, G16, G17, G23, G24 139. Trachinocephalus myops (Forster, 1801) Present study

Crustaceans 1. Penaeus (Penaeus) monodon Fabricius, 1798 G4, G5, G16, G22 2. Penaeus (Penaeus) semisulcatus De Haan, 1844 G5, G16, G23 3. Penaeus (Fenneropenaeus) indicus (H. Milne Edwards, G4, G5, G16, G22, 1837) G23 4. Penaeus (Fenneropenaeus) merguiensis (De Man, 1888) G5, G22, G23 5. Penaeus (Marsupenaeus) japonicus Bate, 1888 G11, G22 6. Metapenaeus dobsoni (Miers, 1878) G4, G5, G8, G16, G18, G22, G23 7. Metapenaeus affinis (H. Milne Edwards, 1837) G4, G5, G8, G16, G22, G23 8. Metapenaeus moyebi (Kishinouye, 1896) G22 9. Parapenaeopsis stylifera (H. Milne Edwards, 1837) G4, G5, G8, G16, G22, G23 10. Parapenaeopsis maxillipedo (Alcock, 1905) Present study 11. Trachypenaeus curvirostris (Stimpson, 1860) G5 12. Exhippolysmata ensirostris (Kemp, 1914) G16 13. Macrobrachium equidens (Dana, 1852) Present study 14. Alpheus sp. G6 15. Panulirus polyphagus (Herbst, 1793) G16 16. Thalassina anomala (Herbst, 1804) Present study 17. Diogenes miles (Fabricius, 1787) Present study 18. Clibanarius infraspinatus Hilgcndorf, 1869 Present study 19. Dorippe astuta (Fabricius, 1793) G6, G13 20. Calappa lophos (Herbst, 1785) Present study 21. Ashtoret lunaris (Forskal, 1775) G6, G13 22. Leucosia pubescens Miers, 1877 G13 23. Doclea rissonii Leach, 1815 G13 24. Portunus (Portunus) sanguinolentus (Herbst, 1783) G6, G13, G16, G18, G23 25. Portunus (Portunus) pelagicus (Linnaeus, 1758) G6, G13, G16, G23 26. Scylla serrata (ForskAl, 1775) G6, G13, G16 27. Scylla olivacea (Herbst, 1796) Present study 28. Charybdis (Charybdis) lucifera (Fabricius, 1798) G13 29. Charybdis (Charybdis) feriata (Linnaeus, 1758) G13,G23 30. Charybdis (Charybdis) variegata (Fabricius, 1798) Present study 31. Charybdis (Charybdis) goaensis Padate et al., 2010 Present study 32. Charybdis (Goniohellenus) vadorum (Alcock, 1899) G13 33. Thalamita crenata (Latreille, 1829) G13 34. Varuna litterata (Fabricius, 1798) G13 35. __ Miyakea nepa (Latreille, 1828) ?? Molluscs 1. Trachycardium f7avum Linnaeus, 1758 G18 2. Dosinia sp. ?? 3. Paphia textile (Gmelin, 1791) G6, G10, G18 4. Anadara granosa (Linnaeus, 1764) G18 5. Solen truncatus Wood, 1815 G18 6. Turritella duplicata (Linnaeus, 1758) G15 7. Bufonaria (Bufonaria) spinosa (Link, 1807) ?? 8. Gyrineum natator (Roding, 1798) G7 9. Murex tribulus Linnaeus, 1758 G15 10. Haustellum (Vokesimurex) malabarim (Smith, 1894) Present study 11. Thais lacera (Born, 1788) G12 12. Babylonia spirata (Linnaeus, 1758) G15 13. Olivancillaria gibbosa (Born, 1778) G15 14. Polinices (Glossaulax) didyma (Roding, 1798) G15 15. Natica pieta Recluz, 1844 G15 16. Natica vitellus vitellus Linnaeus, 1758 G15 17. Cantharus spiralis (Gray, 1839) G15 18. Tibia (Tibia) curta Sowerby, 1842 G15 19. Hydatina velum (Gmelin, 1791) Present study 20. Pugilina cochlidium (Linnaeus, 1758) G15 21. Turricula javana (Linnaeus, 1767) G15 22. Uroteuthis (Photololigo) duvauceli (Orbigny, 1835) G16, G23 23. Sepiella inermis (Van Hasselt, 1835) ?? 24. Cistopus indicus (Orbigny, 1840) Present study

Echinoderms 1. Astropecten indicus Doderlein, 1889 G6 2. Temnopleurus toreumaticus (Leske, 1778) G6

Sea snakes 1. Acrochordus granulatus (Schneider, 1799) G21 2. Enhydrina schistosa (Daudin, 1803) G20, G21 3. Lapemis curtus (Shaw, 1802) G20, G21

Cnidaria 1. Aurelia aurita (Linnaeus, 1758) ??

G1— Rao and Dorairaj (1968) G2 — Talwar (1973) G3 — Tilak (1973) G4 — Prabhu and Dhawan (1974) G5 — George (1980) G6 — Parulekar et aL (1980) G7 — Mookherjee (1985) G8 — Achuthankutty and Parulekar (1986) G9 Poss (1986) G10 — Rao et al. (1992) Gll — Achuthankutty and Nair (1993) G12 — Rao and Rao (1993) G13 — Chatterji (1994) G14 — Ansari et al. (1995) G15 — Apte (1998) G16 — Alvares (2002) G17 — Ansari et al. (2003) G18 — Harkantra and Rodrigues (2003) G19 — Ansari (2004) G20 — Lobo et al. (2004) G21 — Lobo (2005) G22 — Ansari et al. (2006) G23 — Ansari (2008) G24 — Froese and Pauly (2010) ?? — Adequate literature not available Table 3.2. Size class and life stages of species observed during the present study

Faunal group Species Maximum Size of present Life reported size (cm) specimens (cm) stage Stomatopods M. nepa 17.02 7.0 -12.0 J, A Penaeid prawns P. monodon 26.6 (d), 38.0 (Y)2 17.0 - 20.0 J, A P. semisulcatus 18.0 (d), 25.0 (y) 7.0 J P. indicus 18.4 (d), 23.0 (9)2 8.0 -18.0 J, A P. merguiensis 20.0 (d), 24.0 (9)2 11.0 -16.0 J, A P. japonicus 20.0 (d), 30.0 (9)2 9.0 - 15.0 A M. dobsoni 11.8 (d), 13.0 (9)2 6.0 - 9.0 J, A M a.ffinis 14.6 (d), 18.6 (Y?2 8.5 - 9.0 J, A M moyebi 8.3 (d'), 12.6 (9) 5.0 - 7.5 P. stylifera 11.7 (d), 14.5 (Y)2 5.0 - 10.3 J, A P. maxillipedo 10.0 (5), 12.5 (922 2.0 - 5.5 J T. curvirostris 8.1 (d), 10.5 (y) 6.0 Non -'penaeid E. ensirostris 7.92 3.3 - 6.5 J, A prawns M equidens 9.82 6.0 A Lobsters P. polyphagus 40.02 16.0 - 37.0 J, A T. anomala 30.02 22.0 A Anomura D. miles 7.02 4.0 - 6.5 A C. inkaspinatus 8.02 7.0 A Portunid P. sanguinolentus 20.05 4.5 - 14.0 J, A brachyura P. pelagicus 21.05 4.0 - 10.5 J, a S. serrata 28.05 9.0 - 14.8 J, A S. -olivacea 13.85 4.0 -11.2 J, A C. lucifera 10.05 2.5 - 6.0 J, A C. feriata 20.05 3.5 - 7.5 J, a C. variegata 2.05 1.5 - 3.0 A C. goaensis 2.995 2.4 - 3.0 A C. vadorum 2.05 2.5 - 6.0 A T. crenata 8.05 7.0 A Non - portunid D. astuta 1.26 1.0 -1.5 A brachyura C. lophos 10.05 7.0 - 8.0 A A. lunaris 5.05 2.0 - 6.0 A L. pubescens 2.05 1.5 A D. rissonii 5.05 2.2 - 5.0 A V. litterata 5.05 3.0 - 3.5 A Elasmobranchs P. sephen 183.0' 35.0 J N. kuhlii 80.02 14.0 J H. gerrardi 200.02 17.0 J H. walga 45.02 7.5 -18.0 J A. flagellum 72:0' 25.0 - 80.0 J, a R. obtusus 180.0/ 54.0 - 56.0 J G. granulatus 280.02 22.0 - 35.0 J C. griseum 74.02 15.0 - 58.0 J, a S. laticaudus 100.02 15.0 - 58.0 J, a Clupcoid fishes S. longiceps 23.03 10.0 - 20.0 J, A S.brachysoma 13.03 9.5 -15.0 J, A E. thoracata 10.03 8.5 A N. nasus 22.02 9.0 - 21.0 5, A D. acuta 20.03 9.5 - 16.0 J, a 0. tardoore 20.03 7.0 - 14.0 J, a C. dorab 100.03 14.0 - 25.0 J P. ditchela 16.03 7.5 - 13.0 J, A C. dussumieri 20.03 12.0 - 16.0 A 7'. malabarica 17.53 10.5 A T. mystax 15.53 10.0 - 12.0 J, A T. purava 15.5i 10.0 - 22.0 J, a 7'. setirostris 18.03 8.0 -17.0 J, a T dussumieri 11.03 8.5 - 12.0 J, A S. commersonnii 10.02 7.0 -14.0 J, A S. baganensis 10.02 6.5 A Mid catfishes A. maculatus To-.1) 2 3.0 - 22.0 A. subrostratus 39.52 28.0 A N. caelata 45.02 22.0 N. thalassina 185.02 17.0 J Eels M. bagio 200.02 26.0 - 45.0 M. cinereus 220.02 29.0 - 46.0 J P. cancrivorus 108.02 28.0 - 65.0 J Leiognathids G. minuta 21.04 11.5 J P. bindus 11.02 2.0 - 6.5 J, a N. blochii 10.02 6.0 -10.0 J, A L. brevirostris 13.52 8.0 -10.0 J, A L. daura 14.02 7.5 - 14.0 J, A E. splendens 17.02 1.5 - 14.4 J, a S. insidiator 11.33 3.0 -11.5 J, a False trevallies L. lactarius 40.02 3.5 - 15.0 J Threadfin breams N. japonicus 32.02 7.0 -17.0 J, a P. townsendi 20.02 6.5 - 8.5 J Sciaenids D. albida 90.03 21.0 J D. russelii 25.03 12.0 -17.5 J, a Johnius spp. - 30.02 11.0 -19.0 5, a 0. ruber 90.02 10.0 - 23.0 5 P. anea 30.02 2.0 -10.0 5 Ribbon fishes T lepturus 234.02 5.0 - 56.0 J Flatfishes C. arel 40.02 6.0 - 8.0 J C. dubius 50.02 39.0 A C. puncticeps 35.02 13.0 J C. macrostomus 17.32 12.0 -16.5 J, A P. bilineata 30.02 18.0 S. albomaculata 30.02 10.5 -18.5 5, a S. comersonnii 32.02 12.5 - 19.0 5 S. ovata 10.02 4.5 - 7.5 5, A P. triocellatus 15.02 4.0 -10.5 5, a P. arsius 45.02 14.5 - 20.0 J Lizard fishes S. tumbil 60.04 11.0 - 22.0 5, a T. myops 40.02 11.0 J Puffer fishes L. spadiceus 19.91- 6.0 - 25.0 5, A T oblongus 40.02 11.5 J A. immaculatus 30.02 29.0 A C. patoca 38.03 4.5 - 25.0 5, a T. fluviatilis fluviatilis 17.02 12.4 -16.0 A Other teleosts H quoyi 31.23 16.0 -18.0 J 4 H. kuda 30.02 9.5 - 10.0 J M. cephalus 100.03 11.0 - 20.0 J S. canaliculatus 30.02 9.0 - 17.0 J S. argus 38.02 11.0 J A. gymnocephalus 16.02 3.0 - 8.5 J, a A. fasciatus 10.32 8.0 -10.0 A A. bleekeri 10.02 4.0- 7.5 J, a A. indicus 165.02 7.0 - 22.0 J A. djedaba 40.02 10.0 - 20.0 J, a A. atropos 26.52 5.0 - 9.0 J C. malabaricus 60.02 13.5 - 20.5 C. ignobilis 170.02 23.0 J D. russelli 45.02 7.0 -16.5 M. cordyla 80.02 8.0 - 13.0 J P. niger 75.02 2.0 -16.0 S. commersonnianus 120.02 11.0 - 23.0 J S. lysan 110.02 14.0 J R. canadum 200.02 26.0 J H. acuminatus 25.02 4.5 J D. punctata 50.02 16.0 J D. longimana 50.02 9.0 J E. naucrates 110.02 25.0 - 28.0 J G. erythrourus 30.02 9.0 J G. filamentosus 35.02 9.0 -17.0 J G. limbatus 15.02 3.5 - 12.7 J, a G. longirostris 37.02 12.5 - 21.0 J, a L. johnii 97.02 7.5 - 21.0 J L. argentimaculatus 150.02 17.0 C. tuning 60.02 7.5 - 10.0 J M maculata 30.02 8.5 - 12.5 M argenteus 27.03 7.5 J U. vittatus 28.02 13.5 J P. molucca 15.02 6.5 - 8.5 3, a E. tetradactylum 200.02 12.0 J F. heptadactyla 13.03 8.0 -16.0 J, a E. coioides 120.02 11.5 -17.0 J E. diacanthus 55.02 7.0 -17.0 J E. erythrurus 55.02 9.0 - 9.5 J S. sihama 31.02 5.0 - 29.0 J, a A. berda 90.02 16.0 - 29.0 J S. hasta 50.02 27.0 J P. hasta 55.04 11.0 -16.5 J P. furcatus 50.04 14.0 J P. maculatus 59.32 4.0 -12.0 J P. gibbosus 17.03 7.0 - 10.0 J, A P. schotaf 80.02 7.0 - 21.0 J T. jarbua 36.02 10.0 -12.0 J T. theraps 30.02 6.5 - 14.0 3 T. puta 16.02 2.5 - 11.5 J, a S. guttatus 76.04 15.0 - 21.0 J R. kanagurta 35.04 13.0 - 28.0 J, a S. obtusata 55.02 14.0 - 24.5 S. putnamae 90.02 10.0 - 21.0 P. argenteus 60.02 5.5 - 22.0 J P. chinensis 40.02 3.5 - 13.0 J G. giuris 50.03 15.0 5 P. microcephalus 18.02 10.5 A T. vagina 22.02 8.5 - 25.5 5, A O. rubicundus 25.02 6.0 - 14.0 J, a Y. criniger 15.03 10.0 A P.polynema 15.02 6.0 -10.0 J, a O. paulae 6.893 6.5 A C. japonicus 20.03 4.5 - 9.5 5 C. sagitta__ 11.0' 6.0 J E. carbunculus 40.02 7.5 - 12.0 J G. scaber 30.02 14.0 - 21.0 5, a T uranoscopus 8.03 6.5 - 7.5 A T biaculeatus 30.02 3.5 - 20.0 5, a L. cornuta 46.02 2.0 5 C. dussumieri 27.02 10.5 - 18.0 J, A B. mcclellandi 9.63 7.5 A M. gulio 46.02 12.0 J P. lineatus 32.03 7.0 J Cephalopods U. duvauceli 29.07 4.5 - 8.5 J, a S. inermis 12.57 4.5 - 9.5 5, A C. indicus 8.07 6.0 Gastropods T. duplicata 18.08 3.0 - 5.0 J B. spinosa 8.58 2.5 - 3.0 J

G. natator 4.08 1.0 - 1.5 J M tribulus 10.58 4.0 - 4.5 J H. malabaricus 8.58 5.0 - 6.0 A T. lacera 5.58 3.0 - 3.5 B. spirata 7.58 3.0 - 3.5 O.gibbosa 3.08 3.5 - 5.0 P. didyma 9.08 2.0 J N. picta 3.08 1.5 - 2.0 A N. vitellus vitellus 5.08 1.5 - 2.0 J, A C. spiralis 4.08 3.0 - 4.0 A T. curta 18.58 6.0 . 5 H. velum 5.08 2.0 - 3.5 A P. cochlidium 15.08 8.0 A T. javana 7.58 2.0 - 3.0 J Bivalves T,flavum 4.08 1.5 - 2.0 5, A P. textile 8.0' 2.5 5 A. granosa 9.02 2.8 J S. truncatus 5.02 8.0 A Echinoderms A. indicus 4.59 2.0 - 4.0 J, A T toreumaticus 5.010 2.0 J Cnidaria A. aurita 4011 5.0 - 30.0 J, a Sea snakes A. granulatus _ 81.012 67.0 A E. schistosa 140.02 36.0 - 80.0 5, a L. curtus 86.02 45.0 - 50.0 J, a

'Disc width, 2Total length, 3Standard length, 4Fork length, 5Carapace width, 6Carapace length, 7Mantle length, 8Shell height, 9Radius, 1°Horizontal test diameter, "Diameter, 12Snout - vent length, J - population dominated by juveniles, A - adults in large numbers, a - adults rare populations, and adults were occasionally observed (Table 3.2). Further, it was observed that juveniles and young recruits of fin fish and sea snakes dominated the demersal trawl catches during the pre — monsoon, whereas their adults were observed occasionally during the post — monsoon. Among the crustaceans, juveniles and mature adults of penaeid prawns, stomatopods and brachyuran crabs were observed throughout the study duration, whereas adults dominated the non — penaeid prawns and anomuran crabs.

3.3.2. New records for the study area

Out of the two hundred and four taxa collected during the present study, fifty five were recorded for the first time from the region (Table 3.3). These include a newly described portunid crab, Charybdis (Charybdis) goaensis, and a reef — inhabiting teleost, Caesio tuning that was observed for the first time along the entire west coast of India and outside its known bio — geographical range. In addition to these, two crustaceans (Scylla olivacea, C. variegata) and one mollusc (Hydatina velum) are new records for the entire west coast of India. In addition, fifty species including forty two fin fish, six crustaceans (two prawns, one lobster, two hermit crabs, and one brachyura) and two molluscs (one gastropod, and one cephalopod) are recorded for the first time from the region (Table 3.3).

3.3.3. Quantitative analysis

3.3.3a. Overall data

The quantitative analysis of trawl catch data revealed that crustaceans were the most abundant faunal group (57.62 %) during the entire study period (Fig. 3.2a). On the other hand, their contribution was only 45.90 % (Fig. 3.2b) to the total faunal

23 Table 3.3. New records of demersal marine species along Goa coast

Sr. Species Place of Status of report (published) No. occurrence

A. New to science 1. Charybdis (Charybdis) goaensis Padate et al. 2010 I ZM, PFG I This species is new to science2°

B. Extension of geographical range 1. Caesio cuning (Bloch, 1791) PFG First report along the entire west coast of India and outside its known bio — geographical range I9, previously reported only from Port Blair, Andaman islands.5

C. New to West coast of India 1. Scylla olivacea (Herbst, 1796) PFG, MA Pulicat lake' I° 2. Charybdis (Charybdis) variegata (Fabricius, 1798) PFG, ZM, MA ParangipettaiT 3. Hydatina velum (Gmelin, 1791) PFG Southeast coast of India 9

D. New to Goa, central west coast of India 1. Neottygon kuhlii (Muller & Henle, 1841) ZM Maharashtra', Kerala' 2 2. Himantura gerrardi (Gmelin, 1789) PFG Alibaugz Kochi" 3. Himantura walga (Muller & Henle, 1841) PFG, ZM Mumbai 4. Aetobatus flagellum (Bloch & Schneider, 1801) PFG, ZM Kozhikodel8 5. Rhinobatos obtusus Muller & Henle, 1841 PFG Maharashtra, Kerala', Karwar"

6. Thryssa mystax (Bloch & Schneider, 1801) PFG Maharashtra16' 17, Emakulum"

7. Thryssa setirostris (Broussonet, 1782) PFG Maharashtra16' ", Emakulum" 8. Hippocampus kuda Bleeker, 1852 MA Maharashtra16 9. Apogon fasciatus (White, 1870) PFG, ZM South CanaraI3,ahm arashtra16, 17, 18 10. Archamia bleekeri (Gunther, 1859) PFG, ZM Kerala" 11. Scomberoides commersonnianus (Lacepede, 1801) ZM, PFG Maharashtra16' 17, Kerala" 12. Heniochus acuminatus (Linnaeus, 1758) PFG Maharashtra" 13. Drepane longimana (Linnaeus, 1758) ZM, PFG Kozhikode, Emakulum, Kochi" 14. Gerres erythrourus (Bloch, 1791) ZM, PFG Kolaba16, Mumbai, Kochi" 15. Gerres longirostris (Lacepede, 1801) ZM Kerala" 16. Gazza minuta (Bloch, 1795) PFG Mumbai" 17. Leiognathus brevirostris (Valenciennes, 1835) ZIVI Maharashtra16' 17, Mumbai" 18. Monodactylus argenteus (Linnaeus, 1758) MA Kerala, Maharashtra5 19. Parascolopsis townsendi Boulenger, 1901 PFG Kochi18 20. Pempheris molucca Cuvier, 1829 PFG Maharashtra"' ' 7, Mahe" 21. Dendrophysa russelii (Cuvier, 1829) MA Talaja Khadi - Msharashtras 22. Epinephelus coioides (Hamilton, 1822) PFG, ZM Mumbai, Kochi" 23. Epinephelus erythrurus (Valenciennes, 1828) PFG Kerala18 24. Sparidentex hasta (Valenciennes, 1830) MA Kochil8 25. Pomadasys furcatus (Bloch & Schneider, 1801) PFG Kerala' 8 26. Plectorhinchus gibbosus (Lacepede, 1802) MA, ZM Kochi" 27. Plectorhinchus schotaf (Forskal, 1775) ZM, PFG Mumbai, Kerala's 28. Yongeichthys criniger (Valenciennes, 1837) PFG maharashtraik 18 29. Parachaeturichthys polynema (Bleeker, 1853) PFG, MA, ZM Mumbai" 30. Oxyurichthys paulae Pezold, 1998 PFG Kerala" 31. Callionymus japonicus Houttuyn, 1782 PFG, MA, ZM Mangalore 32. Callionymus sagitta Pallas, 1770 PFG, MA Maharashtra16' 17, Kerala' s 33. Eurycephalus carbunculus (Valenciennes, 1833) PFG Mumbai" 34. Synaptura albomaculata Kaup, 1858 PFG, ZM South Canara, Kochi, Kozhikode' 18 35. Triacanthus biaculeatus (Bloch, 1786) MA, ZM, PFG Kasaragod, Cannanore 36. Takifugu oblongus (Bloch, 1786) MA Ratnagiri8, Kolaba", Mumbai, Canara ls 37. Arothron immaculatus (Bloch & Schneider, 1801) PFG Maharashtra"' 17, Kochi" 38. Tetraodonfluviatilis fluviatilis Hamilton, 1822 MA Mumbai, Kozhikode" 39. Arius subrostratus Valenciennes, 1840 MA Kerala, Karnataka' 18 40. Nemapteryx caelata (Valenciennes, 1840) MA Maharashtra16' 17, Mumbai, Kochi" 41. Muraenesox bagio (Hamilton, 1822) ZM Travancore - Kerala18 4

42. Trachinocephalus myops (Forster, 1801) PFG Maharashtra16' 17, Kerala' 43. Parapenaeopsis tnaxillipedo (Alcock, 1905) MA, PFG Mumbai, Kerala(' 44. Macrobrachium equidens (Dana, 1852) MA Kerala& 45. Thalassina anomala (Herbst, 1804) MA Maharashtra3 46. Diogenes miles (Fabricius, 1787) PFG Karwar" 47. Clibanarius infraspinatus Hilgendorf, 1869 ZM Karwar" 48. Calappa lophos (Herbst, 1785) PFG Mumbai, Ratnagiri, Karwar2, Kerala" 49. Haustellum (Vokesimurex) malabaricus (Smith, 1894) ZM Off Malabar coast' 50. Cistopus indicus (Orbigny, 1840) PFG Exact location of report not available

PFG: Potential fishing grounds off Goa MA: Mandovi estuary — Aguada bay region ZM: Zuari estuary — Mormugao bay region

'Alcock (1902) 2Chhapgar (1957) 3Sankolli (1972) 4Joel and Raj (1980) 5Talwar and Kacker (1984) &Kurian and Sebastian (1986) 7Sethuramalingam and Khan (1991) 8 Singh and Yazdani (1993) 9Apte (1998) 1°Keenan et al. (1998) 11Pezold (1998) 12Hanfee (1999) "Mishra and Srinivasan (1999) "Jeyabaskaran et al. (2002) 15Chandran (2003) I6Pathak (2006a) 17Pathak (2006b) "Froese and Pauly (2010) 19 Padate et al. (2010a) 20Padate et al. (2010b) Crustaceans (a) Crustaceans (b) (57.62) (45.90)

Molluscs Molluscs (1.40) (2.52)

Other fauna Other fauna (1.86) (2.11)

Fin fish Fin fish (39.12) (49.46)

Fig. 3.2. Group wise composition of trawl catches observed during the present study (a) numbers and (b) weight weight during the above period. The percentage of fin fish in terms of numbers was only 39.12 % (Fig. 3.2a). However, they dominated the trawl catches in terms of percentage of total weight (49.46 %; Fig. 3.2b). The remaining two groups namely molluscs and "other fauna" contributed only 1.40 and 1.86 % to the total faunal numbers, respectively (Fig. 3.2a). In addition, their contribution to the percentage of total faunal weight during the present study was insignificant (2.52 and 2.11 %, respectively; Fig. 3.2b).

Monthly variations in the numbers and weight of demersal fauna revealed widely fluctuating trends (Fig. 3.3a,b, 3.4a,b). The highest numbers (247,873) and average value (61.97 ± 28.53 x 103) were recorded during March, 2008, whereas the lowest numbers (40,867) and average value (10.22 ± 3.16 x 10 3) were recorded during

February, 2007. In addition, the highest weight (995.45 kg) and average weight

(248.86 ± 167.28 kg) were recorded during March, 2008, whereas the lowest weight

(199.53 kg) and average weight (36.73 ± 17.24 kg) were recorded during April and

May, 2007, respectively. High standard deviation in the average values (Fig. 3.4a,b) indicate large fluctuations in the catches owing to variations among the individual trawl catches.

The temporal trends in the average faunal numbers and weight of demersal fauna computed during the present study revealed generally large fluctuations irrespective of the season (Fig. 3.4a,b). The trawl catches at the commencement of pre

— monsoon, 2006 (February, 2006) were characterized by relatively high numbers

(33.77 ± 4.49 x 103) and weight (146.10 ± 19.48 kg) with relatively large catches of crustaceans (103,664 and 360.41 kg) and fin fish (30,003 and 212.02 kg), followed by large fluctuations up to April, 2006. The average numbers and weight for pre — monsoon, 2006 were 28.64 ± 7.58 x 10 3 and 118.97 ± 30.74 kg, respectively. Further,

24 250 - (a)

0,34 200

150 - k o E 100 z 50

0

1000 - (b) 800 -

600 -

400

200

0 Feb Mar AprDecJan Feb Mar AprMayN ovJan Feb Mar AprMay I 2006 I 2007 I 2008 I

Sampling month

■ Fin fish ■ Crustaceans ■ Molluscs Other fauna

Fig. 3.3. Month wise variations in (a) total numbers and (b) total weight of major faunal groups observed during the present study 90 (a)

3) 10 X ( 60 - bers m nu e

rag 30 Ave

0 i I i 400 - (b)

0 i 1 , 1 1 1 I , 1 , , Feb Mar Apr Dec Jan Feb Mar Apr May Nov Jan Feb Mar Apr May I 2006 I 2007 I 2008 i Sampling month

Fig. 3.4. Month wise variations in (a) average numbers ± S.D. and (b) average weight ± S.D. of demersal fauna observed during the present study the trawl catches during the above period were dominated by "solar prawns"

(Metapenaeus dobsoni and M affinis), and juveniles of most crustacean and fin fish species. Subsequently, in the following post — monsoon season (December, 2006 — January, 2007), there was a sharp decrease in average numbers (14.64 ± 5.17 x 10 3) and marginal reduction in average weight (115.06 ± 39.32 kg). The trawl catches during the above period were characterized by dominance of few coastal species

(Miyakea nepa, Sardinella longiceps and Lagocephalus spadiceus). Thereafter, there was substantial decrease in the average numbers (12.61 ± 5.59 x 10 3) and average weight (48.70 ± 23.24 kg) during pre — monsoon, 2007 with large fluctuations. The above trawl catches were dominated by M dobsoni, Astropecten indicus, and fin fish such as Photopectoralis bindus, Lactarius lactarius and Trichiurus lepturus. The following post — monsoon season (November, 2007 — January, 2008) was characterized by relatively stable trawl catches with substantially higher average numbers (17.44 ± 11.23 x 103), average weight (83.92 ± 35.67 kg), and dominance of

M. nepa, clupeoids and L. spadiceus. Finally, variable trends (sharp fluctuations) were characteristic of trawl catches during pre — monsoon, 2008. The average numbers and average weight during the above period were observed to be 37.16 ± 28.21 x 10 3 and 128.29 ± 111.20 kg, respectively. The only noteworthy observation during pre — monsoon, 2008 was the capture of large shoals of the leiognathid species namely P. bindus during March, 2008. Further computation of temporal trends among the faunal groups revealed the dominance of crustaceans in the individual monthly catches during the present study (Fig. 3.3a,b). However, the fin fish dominated the trawl catches only during February — April, 2007 and March, 2008 (Fig. 3.3a,b). The remaining two groups namely 'other

25 fauna' and molluscs were rare in occurrence and contributed insignificantly to the trawl catches.

Confirmation of the temporal variations in faunal abundance (Table 3.4a,b) and faunal weight (Table 3.5a,b) using two — way analysis of variance (ANOVA) revealed significant differences among the faunal groups between months and seasons at P = 0.001.

3.3.3b. Crustaceans

The "crustaceans", the principal targets of the bottom trawl catches during the present study were divided into seven sub — groups namely penaeid prawns, stomatopods, portunid brachyura, anomura, non — portunid brachyura, non — penaeid prawns and lobsters. Among these, the penaeid prawns dominated the crustacean catches in terms of percentage of numbers and weight, followed by stomatopods and portunid brachyura, whereas the other four sub — groups contributed meagrely (Fig.

3.5a,b). Monthly variations in the crustacean numbers during the present study (Fig.

3.6a, 3.7a) revealed the highest numbers (133,624) and average value (39.81 ± 28.48 x 103) during April, 2008 and May, 2008, respectively, whereas the lowest numbers

(17,025) and average value (4.26 ± 1.66x 10 3) were recorded during February, 2007.

On the other hand, the highest weight (360.41 kg) and average value (90.10 ± 13.04 kg) were recorded during February, 2006 and March, 2008, respectively, whereas, the lowest weight (69.27 kg) and average value (16.25 ± 6.20 kg) were recorded during

February, 2007 and April, 2007, respectively (Fig. 3.6b, 3.7b).

The penaeid prawns (Penaeidae) observed during the present study were represented by eleven species (Metapenaeus dobsoni, M affinis, M moyebi,

Parapenaeopsis stylifera, P. maxillipedo, Penaeus monodon, P. semisulcatus, P.

26 Table 3.4a. Month wise differences in faunal abundance using two - way ANOVA (for months and faunal groups) P = 0.001 Source of Variation SS df MS F P-value F crit Rows 4250.4 26 163.4769 31.5563 2.89E-77 2.16577 Columns 199.3708 14 14.24077 2.748927 0.000693 2.66697 Error 1885.696 364 5.180485 Total 6335.467 404

Table 3.4b. Season wise differences in faunal abundance using two - way ANOVA (for seasons and faunal groups) P = 0.001 Source of Variation SS df MS F P-value F crit Rows 12751.2 26 490.4307 15.44643 4.26E-25 2.392518 Columns 2553.747 4 638.4367 20.10798 2.73E-12 5.000508 Error 3302.043 104 31.75042 Total 18606.99 134

Table 3.5a. Month wise differences in faunal weight using two - way ANOVA (for months and faunal groups) P = 0.001 Source of Variation SS df MS F P-value F crit Rows 8681.766 26 333.9141 27.32762 8.34E-70 2.16577 Columns 526.9065 14 37.63618 3.080156 0.000153 2.66697 Error 4447.687 364 12.21892 Total 13656.36 404

Table 3.5b. Season wise differences in faunal weight using two - way ANOVA (for seasons and faunal groups) P = 0.001 Source of Variation SS df MS F P-value F crit Rows 26045.3 26 1001.742 16.21845 6.32E-26 2.392518 Columns 5453.391 4 1363.348 22.07293 3.28E-13 5.000508 Error 6423.624 104 61.76561 Total 37922.31 134 (a) (b)

Stomatopods Penaeid prawns (31.10) (48.60)

Stomatopods Penaeid prawns (25.77) Anomura (65.75) (0.52)

Portunid brachyura (6.87) Anomura (0.99)

Non - portunid brachyura Portunid brachyura (0.61) (18.76) Non - penaeid prawns Non - penaeid prawns (0.23) (0.48) Non - portunid brachyura (0.33)

Fig. 3.5. Composition of crustaceans in the trawl catches observed during the present study (a) numbers and (b) weight

(a) 140 -

120 -

3) 0 100 1 x

( 80

bers 60

Num 40

20 0

FebMar AprDecJan Feb Mar AprMayNovJan FebMarAprMay I 2006 I 2007 1 2008 I Sampling month ■ Stomatopods ■ Penaeid prawns Non - penaeid prawns ■ Non - portunid brachyura ■ Portunid brachyura ■ Anomura

Fig. 3.6. Month wise variations in (a) total numbers and (b) total weight of crustacean groups observed during the present study Fig. 3.7.Monthwisevariationsin(a)averagenumbers ±S.D.and(b)averageweight Average weight (kg) Average numbers ( x 1 03) 120 40 - 70 - 60 - 90 20 - 30 — 50 - 30 - 60 - 10 - 0 0 1 Feb MarAprDecJanMayNov

± S.D.ofcrustaceansobservedduringthepresent study 2006 I Sampling month 2007 I 2008 (a) (b) 1 indicus, P. merguicnsis, P. japonicus, and Trachypenaeus curvirostris), among which only three species namely M. dobsoni, M. affinis and P. stylifera contributed substantially to the crustaceans in terms of percentage of total numbers (65.75 %; Fig.

3.5a) and weight (48.60 %; Fig. 3.5b). Monthly trends revealed that the penaeid prawns contributed substantially to the crustaceans by total numbers (Fig. 3.6a) and weight (Fig. 3.6b) during the pre — monsoon, particularly during March — April, 2006,

March — May, 2007 and March — May, 2008. During the post — monsoon, their catches were generally lesser than stomatopods. Among the species, M dobsoni and

M. affinis, (euryhaline), were present throughout the present study period, and occurred in comparatively larger quantities during the pre — monsoon. On the other hand, P. stylifera, a purely marine species occurred mostly during the pre — monsoon season. However, during February — May, 2007, this species was obtained in lesser quantities. Other penaeid species (P. monodon, P. indicus, P. merguiensis and P. maxillipedo) occurred in meagre quantities and were inconsistent in their occurrence, thus not making a significant impact on the catch at this time.

The stomatopods represented by Miyakea nepa (Squillidae) during the present study were second among the crustaceans in terms of percentage of total numbers

(25.77 %; Fig. 3.5a) as well as weight (31.10 %; Fig. 3.5b), and were the principal by

— catch of most of the demersal trawl hauls. Monthly trends revealed that M. nepa occurred in relatively high proportions during February, 2006, December, 2006 and

November, 2007 — February, 2008, along with decrease in its catches towards the onset of the southwest monsoon (Fig. 3.6a,b). Further, the above trend was observed to be inversely related to that of the penaeid prawns (Fig. 3.6a,b).

The portunid brachyura (family Portunidae) observed during the present study were represented by ten species (Portunus sanguinolentus, P. pelagicus, Charybdis

27 feriata, C. lucifera, C. goaensis, C. variegata, C. vadorum, Scylla serrata, S. olivacea and Thalamita crenata) those contributed only 6.87 and 18.76 % to the crustaceans by total numbers and weight, respectively (Fig.3.5a,b). This group was represented by juveniles as well as adults, particularly gravid females, throughout the study period.

Monthly trends revealed that the portunid brachyura, particularly the four common species namely P. sanguinolentus, P. pelagicus, C. feriata and C. lucifera occurred regularly in less quantities throughout the present study period (Fig. 3.6a,b). Another species, C. vadorum occurred in few trawls operated at 10 — 20 m depths during

February, April, December, 2006 and January — March, 2007. In addition to these, males and females of C. goaensis were collected from the Zuari estuary in September,

2006 (Padate et al., 2010b), whereas berried females of C. goaensis and S. olivacea, and males of C. variegata were observed occasionally only during the pre — monsoon.

Two other species (S. serrata and T. crenata) were collected only from the estuarine embayment.

The non — portunid brachyura observed during the present study were represented by six species namely Dorippe astuta (Dorippidae), Doclea rissonii

(Majidae), Calappa lophos (Calappidae), Ashtoret lunaris (Matutidae), Leucosia pubescens (Leucosiidae) and Varuna litterata (Grapsidae) those contributed insignificantly to the crustaceans by percentage of total numbers (0.61 %; Fig. 3.5a) and weight (0.33 %; Fig.3.5b). Adult specimens of the above species, except V. litterata, were collected occasionally from soft sandy or muddy substrates those were generally inaccessible to trawlers.

The anomuran crabs (Diogenidae) observed during the present study were represented by two species namely Diogenes miles and Clibanarius infraspinatus, those contributed insignificantly to the crustaceans by percentage of total numbers

28 (0.52 %; Fig.3.5a) and weight (0.99 %; Fig. 3.5b). Monthly trends revealed that the anomuran crabs contributed meagrely to the crustaceans by total numbers (Fig. 3.6a) and weight (Fig. 3.6b) during the present study. D. miles was observed in comparatively larger numbers during the pre — monsoon season, whereas C. infraspinatus (N = 1) was observed once in the Zuary estuary during September,

2007.

The non — penaeid prawns observed during the present study were represented by Exhippolysmata ensirostris (Hippolytidae), Macrobrachium equidens

(Palaemonidae) and Alpheus sp. (Alpheidae) those contributed insignificantly to the crustaceans by percentage of total numbers (0.48 %; Fig. 3.5a) and weight (0.23 %;

Fig. 3.5b). Monthly trends revealed the contribution of the non — penaeid prawns to the total crustacean numbers and weight to be negligible.

The lobsters observed during the present study were represented by Panulirus polyphagus (Palinuridae) and Thalassina anomala (Thalassinidae) those occurred in insignificant numbers in the trawl catches (N = 03 and 01, respectively).

3.3.3c. Fin fish

The "fin fish", the most speciose among the faunal groups observed during the present study were divided into thirteen sub — groups namely elasmobranchs, clupeoid fishes, ariid catfishes, eels, leiognathids, false trevallies, threadfin breams, sciaenids, ribbon fishes, flatfishes, lizard fishes, puffer fishes and other teleosts. Among these, the leiognathids dominated the fin fish catches in terms of percentage of numbers,

followed by clupeoid fishes, other teleosts and puffer fishes (Fig. 3.8a), whereas the other groups contributed meagrely. In terms of fin fish weight, the clupeoid fishes, puffer fishes and leiognathids collectively contributed 60.17 % (Fig. 3.8b), in spite of

29

(a) (b) Elasmobranchs (0.39) Clupeoid fishes (16.06) Other teleosts Other teleosts Elasmobranchs Adid catfishes Puffer fishes (9.09) (9.06) (19.43) (0.78) Puffer fishes (1.62) (7.48) Eels Clupeoid fishes Lizard fishes (0.88) (24.00) (0.64) Lizard fishes Flatfishes (4.95) (2.72) Flatfishes Ribbon fishes (4.27) 7if Ariid catfishes (4.62) Ribbon fishes Sciaenids (1.89) (6.40) (4.83) Sciaenids Eels (8.07) (2.09) Threadfin breams Threadfin bream Leiognathids (0.96) (1.85) (16.74) False trevallies Leiognathids False trevallies (3.31) (47.06) (2.02)

Fig. 3.8. Composition of fin fish in the trawl catches observed during the present study (a) numbers and (b) weight the fact that no single group dominated the catches. Monthly variations in fin fish numbers during the present study (Fig. 3.9a, 3.10a) revealed the highest numbers

(209,862) and average value (52.47 ± 22.13 x 10 3) during March, 2008, whereas the lowest numbers (8,035) and average value (1.61 ± 0.67 x 10 3) were recorded during

November, 2007. In addition, the highest weight (656.80 kg) and average value

(164.20 ± 106.38 kg) were recorded during March, 2008, whereas the lowest weight

(55.29 kg) and average value (12.57 ± 5.10 kg) were recorded during March, 2006 and February, 2008, respectively (Fig. 3.9b, 3.10b).

The elasmobranchs observed during the present study were represented by five rays (Himantura walga, H. gerrardi, Neotrygon kuhlii, Pastinachus sephen

(Dasyatidae) and Aetobatus flagellum (Myliobatidae)), two skates (Rhinobatos obtusus and Glaucostegus granulatus (Rhinobatidae)), and two sharks (Scoliodon laticaudus (Carcharhinidae) and Chiloscyllium griseum (Hemiscyllidae)) those contributed insignificantly to the percentage of fin fish by numbers (0.39 %; Fig. 3.8a) and weight (0.78 %; Fig. 3.8b). Monthly trends revealed that the elasmobranchs contributed insignificantly to the fin fish by total numbers (Fig. 3.9a) and weight (Fig.

3.9b). Among the rays, H. walga juveniles were captured in insignificant quantities (N

< 5 per trawl) in the nearshore trawl hauls almost throughout the study period. In addition, juveniles of the other four rays (H. gerrardi, N. kuhlii, P. sephen and A. flagellum) were observed rarely (N = 1, each) in the nearshore hauls. The skates were rarer in occurrence (N = 7) during the present study and represented only by juveniles mainly during the pre — monsoon season. Among the sharks, C. griseum juveniles were observed to occur commonly in very small numbers throughout the study period, whereas those of S. laticaudus were found mainly during December — February.

30

600 - (b) 500 - Z.; rl.e. 400 - .C" _0) 300 - S

200

Feb Mar Apr Dec Jan Feb Mar Apr May Nov Jan Feb Mar Apr May I 2006 I 2007 2008 Sampling month Elasmobranchs 0 Clupeoid fishes 0 Ariid catfishes Eels ■ ▪ Lelognathids 0 False trevallies Threadfin breams 0 Sciaenids IS Ribbon fishes 0 Flatfishes 0 Lizard fishes Puffer fishes

▪ Other teleosts

Fig. 3.9. Month wise variations in (a) total numbers and (b) total weight of fin fish groups observed during the present study

80 - ( a)

3) 0 1

x 60 - ( rs be

m 40 - nu e rag 20 - Ave

0

300 (b) 250 - C) 200 - a) 150 - a) g 100- 4 50

0 , , I I 1 , I I I I Feb Mar Apr Dec Jan Feb Mar Apr May Nov Jan Feb Mar Apr May I 2006 I 2007 I 2008 I Sampling month

Fig. 3.10. Month wise variations in (a) average numbers ± S.D. and (b) average weight ± S.D. of fin fish observed during the present study Adults of both these species (N = 1, each) were observed only on one occasion each during the post — monsoon season.

The clupeoid fishes observed during the present study were represented by sixteen species namely Sardinella longiceps, S. brachysoma, Escualosa thoracata,

Nematalosa nasus, Dussumieria acuta, Opisthopterus tardoore (Clupeidae), Pellona ditchela (Pristigasteridae), Coilia dussumieri, Stolephorus commersonnii, S. baganensis, Thiyssa dussumieri, T. mystax, T. malabarica, T. purava, T. setirostris

(Engraulidae) and Chirocentrus dorab (Chirocentridae) those contributed substantially to the fin fish by percentage of total numbers (16.06 %; Fig. 3.8a) and weight (24.00 %; Fig. 3.8b). Monthly trends revealed that clupeoid fishes contributed substantially to the fin fish by total numbers (Fig. 3.9a) and weight (Fig. 3.9b).

Among the species, S. longiceps was observed throughout the present study (except

March and May of each sampling year) and its numbers generally peaked during

November — January. Juveniles and adults of the anchovies (Family Engraulidae) were observed throughout the present study. Other clupeoid species were either observed in lesser quantity throughout the present study (0. tardoore and P. ditchela) or occasionally (S. brachysoma, D. acuta, N. nasus and C. dorab) mostly as juveniles.

In addition, adults of some species (E. thoracata, N. nasus and D. acuta) were observed in insignificant numbers in estuarine samples during the monsoon season.

The ariid catfishes (Family Ariidae) observed during the present study were represented by four species namely Arius maculatus, A. subrostratus, Nemapteryx caelata and Netuma thalassina (comprising mostly juveniles) those contributed insignificantly to the fin fish by percentage of total numbers (1.52 %; Fig. 3.8a) and weight (1.89 %; Fig. 3.8b). Monthly trends revealed that the ariid catfishes contributed meagrely to the fin fish by total numbers (Fig. 3.9a) and weight (Fig.

31 3.9b). The dominant species namely A. maculatus was observed regularly in small numbers in the nearshore bottom trawls during the entire study period. The other three species were obtained occasionally during sampling surveys in the adjacent bay — estuarine waters.

The eels observed during the present study comprised exclusively juveniles of three species namely Muraenesox bagio, M cinereus (Muraenesocidae) and

Pisodonophis cancrivorus (Ophichthidae) those contributed insignificantly to the fin fish by percentage of total numbers (0.88 %; Fig. 3.8a) and weight (2.09 %; Fig.

3.8b). Monthly trends revealed that the eels contributed meagrely to the fin fish by total numbers (Fig. 3.9a) and weight (Fig. 3.9b). Among the species, few individuals of M cinereus (N < 5) were observed in majority of the nearshore trawls during

December to April of each sampling year. On the other hand, M bagio was caught only once from the Zuari estuary and P. cancrivorus was observed occasionally in nearshore trawls during the pre — monsoon season.

The leiognathids (Family Leiognathidae) observed during the present study were represented by seven species namely Gazza minuta, Leiognathus brevirostris, L. daura, Nuchequula blochii, Photopectoralis bindus, Eubleekeria splendens and

Secutor insidiator those contributed substantially to the fin fish by percentage of total numbers (47.05 %; Fig. 3.8a) and weight (16.74 %; Fig. 3.8b). Most of the above species (except P. bindus and G. minuta) were represented by juveniles as well as adults. Monthly trends revealed that the leiognathids contributed sizeably to the fin fish by total numbers (Fig. 3.9a) and weight (Fig. 3.9b) with wide variations. Among the species, P. bindus was the most abundant and alone accounted for 44.27 and 14.99

% of the fin fish by total numbers and weight, respectively. Monthly trends revealed that this species occurred during most of the sampling months with a peak during

32 April, 2006, March — May, 2007 and March — April, 2008. Large shoals of P. bindus juveniles (2.0 — 6.0 cm TL) weighing upto 169 kg were caught during March, 2008.

In addition, two other species namely E. splendens and S. insidiator were observed in low numbers in most hauls during the present study, whereas the other four species

(G. minuta, L. brevirostris, L. daura and N. blochii) were found occasionally in insignificant numbers.

The false trevallies (Family Lactariidae) observed during the present study were represented exclusively by Lactarius lactarius juveniles those contributed insignificantly to the fin fish by percentage of total numbers (3.31%; Fig. 3.8a) and weight (2.02 %; Fig. 3.8b). Monthly trends revealed that the false trevallies contributed meagrely to the fin fish by total numbers (Fig. 3.9a) and weight (Fig.

3 .9b).

The threadfin breams (Family Nemipteridae) observed during the present study were represented by two species namely Nemipterus japonicus and

Parascolopsis townsendi those contributed insignificantly to the fin fish by percentage of total numbers (0.96 %; Fig. 3.8a) and weight (1.85 %; Fig. 3.8b). Among these, N. japonicus was mostly represented by juveniles and occasionally by adults, whereas P. townsendi was represented only by juveniles. Monthly trends revealed that the threadfin breams contributed meagrely to the fin fish by total numbers (Fig. 3.9a) and weight (Fig. 3.9b). Among the species, N. japonicus appeared in very low numbers with a peak in December, 2006 and subsequent reduction up to March, 2007. There was a second peak in March, 2008 with comparatively lower numbers. In addition, juveniles of the other species (P. townsendi) were observed occasionally during the pre — monsoon season.

33 The sciaenids (Family Sciaenidae) observed during the present study comprised mostly juveniles of four species namely Daysciaena albida, Dendrophysa russelii, Otolithes ruber, Pennahia anea, and unidentified Johnius spp. those contributed meagrely to the fin fish by percentage of total numbers (5.40 %; Fig. 3.8a) and weight (8.07 %; Fig. 3.8b). Monthly trends revealed that the sciaenids occurred regularly in trawl hauls, however contributed meagrely to the fin fish by total numbers

(Fig. 3.9a) and weight (Fig. 3.9b).

The ribbon fishes (Family Trichiuridae) observed during the present study were represented mostly by juveniles of Trichiurus lepturus those contributed meagrely to the fin fish by percentage of total numbers (4.52 %; Fig. 3.8a) and weight

(4.83 %; Fig. 3.8b). Monthly trends revealed that although the ribbon fishes occurred regularly in trawl hauls, they contributed meagrely to the fin fish by total numbers

(Fig. 3.9a) and weight (Fig. 3.9b). Adults of T. lepturus were observed only during

November, 2007.

The flatfishes observed during the present study were represented by ten species namely Cynoglossus arel, C. dubius, C. puncticeps, C. macrostomus,

Paraplagusia bilineata (Cynoglossidae), Solea ovata, Synaptura albomaculata, S. commersonnii (Soleidae), Pseudorhombus triocellatus and P. arsius (Paralichthyidae) those contributed meagrely to the fin fish by percentage of total numbers (2.72 %;

Fig. 3.8a) and weight (4.27 %; Fig. 3.8b). Monthly trends revealed that the flatfishes contributed meagrely to the fin fish by total numbers (Fig. 3.9a) and weight (Fig.

3.9b). The most common flatfish namely C. macrostomus occurred almost throughout the study, except May, 2008, whereas the other cynoglossids occurred rarely. Among the soles, S. ovata adults occurred in insignificant numbers occasionally throughout the present study. Among the flounders, P. triocellatus juveniles occurred commonly

34 in meagre numbers, mostly during the pre — monsoon, whereas P. arsius occurred rarely.

The lizard fishes (Family Synodontidae) observed during the present study comprised mostly juveniles of two species namely Saurida tumbil and

Trachinocephalus myops those contributed insignificantly to the fin fish by percentage of total numbers (0.64 %; Fig. 3.8a) and weight (4.95 %; Fig. 3.8b).

Monthly trends revealed that the lizard fishes occurred only during December, 2006 —

March, 2007 and January and March, 2008 and contributed meagrely to the fin fish by total numbers (Fig. 3.9a) and weight (Fig. 3.9b). Among the species, S. tumbil constituted the bulk of the lizard fish catch, whereas T. myops (N = 1) appeared only once during the present study. Adults of S. tumbil were observed mostly during

January, 2007 and 2008.

The puffer fishes (Family Tetraodontidae) observed during the present study were represented by four species namely Lagocephalus spadiceus, Chelonodon patoca, Arothron immaculatus, Takifugu oblongus, Tetraodon (Tetraodon)fluviatilis, and contributed meagrely to the fin fish by percentage of total numbers (7.48 %; Fig.

3.8a) and weight (19.43 %; Fig. 3.8b). Monthly trends revealed that the puffer fishes contributed substantially to the fin fish by total numbers (Fig. 3.9a) and weight (Fig.

3.9b). Among the species, L. spadiceus contributed to the bulk of puffer fish catches with peaks in numbers during February, 2006, January — February, 2007 and January

— February, 2008. In addition, the other marine puffers (A. immaculatus, 7'. oblongus,

N = 1 each, and C. patoca, N = 6) were observed rarely during the entire study, whereas the freshwater T fluviatilis fluviatilis was obtained (N = 4) only from the estuarine surveys during the southwest monsoon season.

35 The "other teleosts" observed during the present study constituted the remainder of fin fish those were not included in any of the above twelve sub — groups, and was the most diverse taxonomic group comprising seventy four species from thirty eight families belonging to nine orders (Table 3.6). This sub — group contributed only 9.05 and 9.09 % to the total fin fish by numbers and weight, respectively (Fig. 3.8a,b). Monthly trends revealed that this sub — group contributed meagrely to the fin fish by total numbers (Fig. 3.9a) and weight (Fig. 3.9b). Among these species, only P. lineatus showed typical shoaling feature during March, 2006 and April, 2007 wherein large number of juveniles of almost uniform length (7 cm

TL) were observed in three separate trawl hauls at 5 — 13 m depths.

3.3.3d. Molluscs

The "molluscs", one of the minor by — catch groups observed during the present study were divided into three sub —groups namely cephalopods, gastropods and bivalves. Among these, the cephalopods dominated the molluscs in terms of percentage of total numbers as well as weight, whereas the other two groups contributed insignificantly (Fig. 3.11a,b). Monthly variations in the numbers of molluscs during the present study (Fig. 3.12a, 3.13a) revealed the highest numbers

(7,586) and average value (1.26 ± 1.07 x 10 3) during April, 2008, whereas the lowest numbers (202) and average value (0.04 ± 0.07 x 10 3) were recorded during

November, 2007. In addition, the highest weight (41.56 kg) and average value (10.36

±10.07 kg) were recorded during April, 2008 and March, 2008, respectively, whereas the lowest weight (1.60 kg) and average value (0.32 ± 0.37 kg) were recorded during

April, 2007 and November, 2007, respectively (Fig. 3.12b, 3.13b).

36 Table 3.6. List of species categorized as "other teleosts" within the fin fish component of the demersal community during the present study

Sr. Fin fish orders Constituent species No. (family name in parenthesis) 1. Beloniformes Hyporhamphus quoyi (Hemiramphidae) 2. Syngnathiformes Hippocampus kuda (Syngnathidae) 3. Mugiliformes Mugil cephalus (Mugilidae) 4. Perciformes Siganus canaliculatus (Siganidae), Scatophagus argus (Scatophagidae), Ambassis gymnocephalus (Ambassidae), Apogon fasciatus, Archamia bleekeri (Ambassidae), Alectis indicus, Alepes djedaba, Atropus atropos, Carangoides malabaricus, Caranx ignobilis, Decapterus russelli, Megalaspis cordyla, Parastromateus niger, Scomberoides commersonnianus, S. lysan (Carangidae), Rachycentron canadum (Rachycentridae), Heniochus acuminatus (Chaetodontidae), Drepane longimana, D. punctata (Drepaneidae), Echeneis naucrates (Echeneidae), Gerres erythourus, G. filamentosus, G. limbatus, G. longirostris (Gerreidae), Lutjanus johnii, L. argentimaculatus (Lutjanidae), Caesio cuning (Caesionidae), Mene maculata (Menidae), Monodactylus argenteus (Monodactylidae), Upeneus vittatus (Mullidae), Pempheris molucca (Pempheridae), Eleutheronema tetradactylum, Filimanus heptadactyla (Polynemidae), Epinephelus diacanthus, E. coioides, E. erythrurus (Serranidae), Sillago sihama (Sillagindae), Acanthopagrus berda, Sparidentex hasta (Sparidae), Pomadasys hasta, P. furcatus, P. maculatus, Plectorhinchus gibbosus, P. schotaf(Haemulidae), Terapon jarbua, T puta, T theraps (Terapontidae), Rastrelliger kanagurta, Scomberomorus guttatus (Scombridae), Sphyraena obtusata, S. putnamae (Sphyraenidae), Pampus argenteus, P. chinensis (Stromateidae), Glossogobius giuris, Paratrypauchen microcephalus, Trypauchen vagina, Odontamblyopus rubicundus, Yongeichthys criniger, Parachaeturichthys polynema, Oxyurichthys paulae (Gobiidae), Callionymus japonicus, C. sagitta (Callionymidae) 5. Scorpaeniformes Grammoplites scaber, Eurycephalus carbunculus (Platycephalidae), Trachicephalus uranoscopus (Synanceiidae) 6. Tetraodontiformes Triacanthus biaculeatus (Triacanthidae), Lactoria cornuta (Ostracioniidae) 7. Batrachoidiformes Colletteichthys dussumieri (Batrachoididae) 8. Gadiformes Bregmaceros mcclellandi (Bregmacerotidae) 9. Siluriformes Mystus gulio (Bagridae), Plotosus lineatus (Plotosidae) (a) (b)

Gastropods Bivalves Gastropods Bivalves 1.84) (0.19) (7.29) (2.18)

Cephalopods Cephalopods (90.87) (97.63)

Fig. 3.11. Composition of molluscs in the trawl catches observed during the present study (a) numbers and (b) weight

(a)

0

40 (b)

30 -

10 1 7

Feb Mar Apr Dec Jan Feb Mar Apr May Nov Jan Feb Mar Apr May I 2006 I 2007 I 2008 Sampling month

[I Cephalopods Gastropods ❑ Bivalves

Fig. 3.12. Month wise variations in (a) total numbers and (b) total weight of mollusc groups observed during the present study 3- (a)

3) 10 x (

rs 2- be m nu e 1 Averag

0 4...., Alliiiiiiik rii111111b.. -moo

20 - (b)

0 Feb Mar Apr Dec Jan Feb Mar Apr May Nov Jan Feb Mar Apr 1------2006-----I------2007------I------2008------1

Sampling month

Fig. 3.13. Month wise variations in (a) average numbers ± S.D. and (b) average weight ± S.D. of molluscs observed during the present study The cephalopods observed during the present study were represented by three species namely Uroteuthis (Photololigo) duvauceli (Loliginidae), Sepiella inermis

(Sepiidae) and Cistopus indicus (Octopodidae) those dominated the catches of molluscs in terms of percentage of total numbers (90.87 %; Fig. 3.11a) and weight (97.63 %; Fig. 3.11b). Monthly trends revealed the dominance of the cephalopods with peaks in numbers during March, 2006, December, 2006, May, 2007 and February — May, 2008 (Fig. 3.12a,b). The gastropods observed during the present study were represented by sixteen species namely Turritella duplicata (Turritellidae), Bufonaria (Bufonaria) spinosa

(Bursidae), Gyrineum natator (Ranellidae), Tibia (Tibia) curta (Strombidae), Murex tribulus, Haustellum (Vokesimurex) malabaricus, Thais lacera (Muricidae),

Babylonia spirata (Babyloniidae), Olivancillaria gibbosa (Olividae), Natica pitta, N. vitellus vitellus, Polinices (Glossaulax) didyma (Naticidae), Cantharus spiralis

(Buccinidae), Pugilina cochlidium (Melongenidae), Turricula javana (Clavatulidae) and Hydatina velum (Aplustridae) those contributed only 7.29 and 2.18 % to the total numbers and weight of molluscs, respectively (Fig. 3.11a,b.) Monthly trends revealed insignificant quantities of these species (Fig. 3.12a,b). The bivalves observed during the present study were represented by

Trachycardium flavum (Cardiidae), Dosinia sp., Paphia textile (Veneridae), Solen truncatus (Solenidae) and Anadara granosa (Arcidae) those contributed only 1.84 and 0.19 % to the total numbers and weight of molluscs, respectively (Fig. 3.11a,b.) Monthly trends revealed insignificant quantities of these species (Fig. 3.12a,b).

37 3.3.3e. Other fauna

The "other fauna" observed during the present study were divided into four sub — groups namely echinoderms, cnidaria, sea snakes and miscellaneous (juveniles and unidentified species). Among these, the echinoderms were the highest in terms of percentage of total numbers (78.27 %; Fig. 3.14a), whereas the cnidaria dominated by percentage of total weight (52.92 %; Fig. 3.14b). Monthly variations in the numbers of the other fauna during the present study (Fig. 3.15a, 3.16a) revealed the highest numbers (9,406) and average value (2.05 ± 1.30 x 10 3) during May, 2007 and March,

2008, respectively, whereas the lowest values (zero) were recorded during December,

2006, and January — February, 2008. In addition, the highest weight (45.43 kg) and average value (11.36 ± 21.72 kg) were recorded during March, 2008, whereas the lowest values (zero) were recorded during January, 2008 (Fig. 3.15b, 3.16b).

The echinoderms observed during the present study were represented by two species namely Astropecten indicus (Astropectinidae) and Temnopleurus toreumaticus (Temnopleuridae) those dominated the catches of other fauna in terms of percentage of total numbers (78.27 %; Fig. 3.14a), however, they contributed only

34.00 % to the total other fauna weight (Fig. 3.14b). Monthly trends revealed the dominance of the echinoderms during March — April, 2006, February — May, 2007 and April — May, 2008 (Fig. 3.15a,b). Among the species, A. indicus was observed in higher quantities during the pre — monsoon with peak numbers (9,335) and weight

(16.03 kg) during May, 2007. The other species namely T. toreumaticus (N = 2) occurred once during the study.

The cnidaria represented by Aurelia aurita (Family Ulmaridae) during the present study, were second among the other fauna in terms of percentage of total numbers (17.41 %; Fig. 3.14a) and highest in terms of percentage of total weight

38 (a) (b)

Miscellaneous Miscellaneous (3.75) (0.45) Echinoderms (34.00) Cnidaria (17.41 Sea snakes (0.57)

Cnidaria (52.92)

Echinoderms Sea snakes (78.27) (12.64)

Fig. 3.14. Composition of other fauna in the trawl catches observed during the present study (a) numbers and (b) weight 10 - (a)

3) 8 10

x ( rs be m Nu

0 50 -

40

30

20

10-

0 --I. Feb Mar AprDec Jan Feb Mar Apr MayN ovJan Feb Mar Apr May 2006 I 2007 I 2008 I Sampling month o Echinoderms Sea snakes

o Cnidaria ❑ Miscellaneous

Fig. 3.15. Month wise variations in (a) total numbers and (b) total weight of other fauna groups observed during the present study 3- (a)

3) 10 x

( 2- rs be m nu e rag Ave

30- (b)

as 20 a)

cs) co 436" 10 - >

Feb Mar Apr Dec Jan Feb Mar Apr May Nov Jan Feb Mar Apr May I Sampling month

Fig. 3.16. Month wise variations in (a) average numbers ± S.D. and (b) average weight ± S.D. of other fauna observed during the present study (52.92 %; Fig. 3.14b). Monthly trends revealed marked presence of A. aurita in the trawl catches during March, 2006, November, 2007 and March, 2008 (Fig. 3.15a,b) with peak numbers (4,057) and weight (45.43 kg) during March, 2008.

The sea snakes observed during the present study were represented by three species namely Enhydrina schistosa, Lapemis curtus (Hydrophiidae) and

Acrochordus granulatus (Acrochordidae), and although occuring regularly in the trawl catches, they contributed insignificantly to the other fauna in terms of percentage of total numbers (0.57 %; Fig. 3.14a) and weight (12.64 %; Fig. 3.14b).

Monthly trends revealed occasional peaks in their numbers during April, 2006 and

February, 2007 (Fig. 3.15a; Padate et al., 2009).

The miscellaneous group observed during the present study comprised unidentified juveniles those contributed insignificantly to the other fauna in terms of percentage of total numbers (3.75 %; Fig. 3.14a) and weight (0.45 %; Fig. 3.14b).

Monthly trends revealed insignificant numbers with comparatively high numbers (446 and 557) during February, 2006 and January, 2007, respectively.

3.3.3f Coastal residents and migrants

Among the two hundred and four demersal taxa observed during the present study, only one hundred and eighteen were observed regularly in the bottom trawl hauls. Further, sixty nine among these were grouped as 'Coastal residents' and the remaining forty nine as 'Migrants' (Froese and Pauly, 2010; Palomares and Pauly,

2010). The coastal residents comprised mostly adults and juveniles of crustaceans, molluscs, echinoderms, cnidaria and some fin fish (Table 3.7). On the other hand, the migrant species comprised mostly fin fish juveniles (Table 3.7). Quantitative analysis revealed that the coastal residents were dominant in the study area and contributed

39 Table 3.7. Habitat use guilds of commonly observed demersal species during the present study

Habitat use guild Constituent taxa Coastal residents Himantura walga, Sardinella sp., Dussumieria acuta, Thryssa setirostris, Archamia bleekeri, Alectis indicus, Decapterus russelli, Megalaspis cordyla, Mene maculata, Siganus canaliculatus, Leiognathus daura, Lactarius lactarius, Nemipterus japonicus, Sphyraena obtusata, Sphyraena putnamae, Filimanus heptadactyla, Pennahia anea, Epinephelus diacanthus, Rastrelliger kanagurta, Pempheris molucca, Callionymus japonicus, Oxyurichthys paulae, Paratrypauchen microcephalus, Parachaeturichthys polynema, Cynoglossus macrostomus, C. puncticeps, Solea ovata, Pseudorhombus triocellatus, Trachinocephalus myops, Triacanthus biaculeatus, Colletteichthys dussumieri, Miyakea nepa, Penaeus indicus, P. merguiensis, Penaeus sp., Metapenaeus dobsoni, M. affinis, Metapenaeus sp., Parapenaeopsis stylifera, P. maxillipedo, Parapenaeopsis sp., Trachypenaeus curvirostris, Exhippolysmata ensirostris, Alpheus sp., Dorippe astuta, Doclea rissoni, Calappa lophos, Ashtoret lunaris, Portunus sanguinolentus, P. pelagicus, Charybdis feriata, C. lucifera, C. goaensis, C. variegate, C. vadorum, Unidentified xanthid crab, Diogenes miles, Clibanarius infraspinatus, Uroteuthis (Photololigo) duvauceli, Sepiella inermis, Cistopus indicus, Turritella sp., Bufonaria sp., Unidentified gastropod, Anadara sp., Astropecten indicus, Lapemis curtus, Enhydrina schistosa, Aurelia aurita Migrants a. Amphidromous Scoliodon laticaudus, Opisthopterus tardoore, Chirocentrus dorab, Coilia dussumieri, Stolephorus baganensis, Thryssa dussumieri, Plotosus lineatus, Grammoplites scaber, Ambassis gymnocephalus, Alepes djedaba, Atropus atropos, Parastromateus niger, Pampus chinensis, Terapon puts, Photopectoralis bindus, Leiognathus brevirostris, Nuchequula blochii, Eubleekeria splendens, Secutor . insidiator, Gerres filamentosus, G. limbatus, Dendrophysa russelli, Johnius spp., Otolithes ruber, Odontamblyopus rubicundus, Trypauchen vagina, Saurida tumbil b. Anadromous Nematalosa nasus, Pellona ditchela, Stolephorus commersonnii, Thryssa mystax, Pisodonophis cancrivorus c. Catadromous Terapon jarbua d. Marine migrant Arius maculatus, Muraenesox bagio, Mugil cephalus, Trichiurus lepturus, e. Marine straggler Chiloscyllium griseum, Sardinella longiceps, Thryssa purava, Muraenesox cinereus, Apogon fasciatus, Pampus argenteus, Pomadasys maculatus, Scomberomorus guttatus, Synaptura albomaculata, Lagocephalus spadiceus, Bregmaceros mcclellandi f. Estuarine Sillago sihama migrant 56.76 and 64.69 % to the total faunal numbers and weight, respectively. Monthly trends in their numbers revealed their dominance in most of the nearshore trawl catches during the present study, except April, 2007 and March, 2008, whereas in terms of weight they dominated throughout the study, except April, 2007 and March, 2008 (Fig. 3.17).

3.3.3g. Diversity The present values for species diversity, evenness and richness ranged between 1.03 (March, 2008) and 2.73 (February, 2007); 0.00 (March, 2008) and 0.09 (November, 2007) and 0.62 (December, 2006) and 3.90 (May, 2007), respectively Table 3.8; Fig. 3.18). Inversely, dominance values were found to be exceptionally high during March, 2008 (0.59; Fig. 3.18). Conversely, lowest dominance values coincided with highest diversity and evenness. Monthly variations in the faunal diversity, evenness and richness during 2006, 2007 were characterized by an increasing trend with the progress of the post — monsoon season (November — January), followed by a peak during February — March and subsequent reduction towards the late pre — monsoon (April — May; Fig. 3.18). However, the present observations revealed large fluctuations in the diversity indices during pre — monsoon, 2008 (February — May, 2008). One — way ANOVA indicated significant monthly variation in the diversity indices (Table 3.9a — d) at P = 0.001 level of significance. On the other hand, no significant variations were observed between sampling seasons (Table 3.10a — d) even at P = 0.05 level of significance.

40 80 - A A Coastal resident species (a)

3) e--o Migrant species 10

x GO - ( rs be m 40 - nu e rag Ave

(b) 200 - Si

0, 150 cr) 100

50 '''411111k

0 1 1 1 1 1 1 1 1 I Feb Mar Apr Dec Jan Feb Mar Apr May Nov Jan Feb Mar Apr May I— 2006---I 2007------1------2008----1

Sampling month

Fig. 3.17. Month wise trends in (a) average numbers ± S.D. and (b) average weight ± S.D. of coastal residents and migrant species observed during the present study Table 3.8. Range and averages of species diversity indices computed during the present study

Sr. No. Diversity index Range of values Mean ± S.D. 1. Shannon — Wiener' diversity index 1.03 — 2.73 1.94 ± 0.44 2. Heip's evenness index 0.00 — 0.09 0.05 ± 0.02 3. Margalef s species richness 0.62 — 3.90 2.04 ± 0.66 4. Simpson's dominance index 0.10 — 0.59 0.27 ± 0.12 Feb Mar Apr Deo Jan Feb Mar Apr Mayne, Jan Fell Mar Apr May

Sarrpling month

Fig. 3.18. Month wise variations in species diversity indices (average f S.D.) of the demersal fish community during the present study Table 3.9a. Month wise differences in diversity using one - way ANOVA (P = 0.001) P = 0.001 Source of Variation SS df MS F P-value F crit

Between Groups 8.338974 14 0.595641 6.611813 1.78E- 07 3.229817 Within Groups 4.774632 53 0.090087 Total 13.11361 67

Table 3.9b. Month wise differences in evenness using one - way ANOVA (P = 0.001) P = 0.001 Source of Variation SS df MS F P-value F crit

Between Groups 0.010962 14 0.000783 4.305877 5.05E-05 3.229817 Within Groups 0.009638 53 0.000182 Total 0.0206 67

Table 3.9c. Month wise differences in species richness using one - way ANOVA (P = 0.001) P = 0.001 Source of Variation SS df MS F P-value F crit

Between Groups 15.64869 14 1.117764 4.509975 2.94E -05 3.229817 Within Groups 13.13565 53 0.247843 Total 28.78435 67 Table 3.9d. Month wise differences in dominance using one - way ANOVA (P = 0.001)

P = 0.001 Source of Variation SS df MS F P-value F crit

Between Groups 0.662436 14 0.047317 7.427635 3.03E - 08 3.229817 Within Groups 0.33763 53 0.00637 Total 1.000066 67 Table 3.10a. Season wise differences in diversity using one - way ANOVA (P = 0.05) P = 0.05 Source of Variation SS df MS F P-value F crit Between Groups 0.096309 4 0.024077 0.116527 0.976197 2.51767 Within Groups 13.0173 63 0.206624 Total 13.11361 67

Table 3.10b. Season wise differences in evenness using one - way ANOVA (P = 0.05) P = 0.05 Source of Variation SS df MS F P-value F crit Between Groups 0.001292 4 0.000323 1.054005 0.386755 2.51767 Within Groups 0.019308 63 0.000306 Total 0.0206 67

Table 3.10c. Season wise differences in species richness using one - way ANOVA (P = 0.05) P = 0.05 Source of Variation SS df MS F P-value F crit Between Groups 2.089967 4 0.522492 1.233105 0.305927 2.51767 Within Groups 26.69438 63 0.42372 Total 28.78435 67

Table 3.10d. Season wise differences in dominance using one - way ANOVA (P = 0.05) P = 0.05 Source of Variation SS df MS F P-value F crit Between Groups 0.011979 4 0.002995 0.190939 0.942279 2.51767 Within Groups 0.988087 63 0.015684 Total 1.000066 67 3.3.4. Faunal Associations

Associations among demersal marine fauna observed during the tenure of present study (Table 3.11) were explained on the basis of four categories of ecological guilds (Elliot et al., 2007; Table 3.12).

3.3.4a. Family level

The cluster analysis of numerical data of fish families collected during the present study revealed altogether twenty four clusters with > 50 % similarity (Fig.

3.19a,b). Among these, only three clusters were selected to investigate and examine in order to generate comprehensive information on the different types of associations among these taxa. Cluster I comprised the three most abundant families namely

Penaeidae, Leiognathidae and Squillidae. Among these, Penaeidae and Squillidae showed 80 % similarity among themselves, and both these clustered with

Leiognathidae at 70 % similarity. Cluster V comprised fifteen families, among which three sub — clusters, namely a) Clupeidae, Engraulidae, Sciaenidae and

Trichiuridae, b) Tetraodontidae and Port-unidae, and c) Lactariidae, Cynoglossidae and Sepiidae showed a similarity of > 80 % whereas, other three sub — clusters namely a) Diogenidae and Loliginidae, b) Muraenesocidae and Haemulidae, and c)

Ariidae and Astropectinidae showed a similarity of < 80 %. Cluster XIX comprised

Nemipteridae and Synodontidae with 80 % similarity among them.

3.3.4b. Family level: pre — monsoon season

The cluster analysis of numerical data of fish families for pre — monsoon months collected during the present study revealed altogether twenty clusters with >

50 % similarity (Fig. 3.20a,b). Among these, only three clusters were selected to

41 Table 3.11. Ecological criteria observed for the major fish families during the present study

Sr. Family Habitat Migration Diet preference No. 1. Clupeidae Pelagic, up to 200 Migrants Phytoplankton, m depth (amphidromous, zooplankton oceanodromous) 2. Engraulidae Bentho — pelagic, Migrants Phytoplankton, up to 50 m depth (amphidromous, zooplankton, teleosts, oceanodromous) prawns 3. Sciaenidae Bentho — pelagic, Migrants Teleosts, prawns and up to 50 m depth (amphidromous) benthic invertebrates 4. Trichiuridae Bentho — pelagic, Migrants Teleosts, prawns, up to 350 m depth (amphidromous) benthic invertebrates, zooplankton 5. Portunidae Demersal, up to 30 Non — migrants Benthic invertebrates, m depth detritus 6. Tetraodontidae Demersal, coastal Migrants Teleosts, cephalopods, waters (varied) crabs 7. Lactariidae Pelagic, up to 100 Non — migrants Teleosts, zooplankton, m benthic invertebrates 8. Leiognathidae Demersal, up to Migrants Phytoplankton, 100 m (amphidromous) zooplankton, benthic invertebrates 9. Squillidae Demersal, up to 40 Non — migrants Benthic invertebrates m depth 10. Penaeidae Demersal, up to 90 Non — migrants Plankton, benthic m depth invertebrates, algae, detritus 11. Cynoglossidae Demersal, Non — migrants Benthic invertebrates, commonly up to 25 some feed on plankton m depth and small teleosts 12. Muraenesocidae Demersal, up to Migrants Teleosts, prawns, other 740 m depth (oceanodromous) benthic crustaceans 13. Haemulidae Demersal, up to Migrants Teleosts, benthic 100 m depth (amphidromous) invertebrates 14. Ariidae Demersal, up to Migrants Teleosts, algae, and 200 m depth (varied) variety of benthic invertebrates 15. Sepiidae Demersal, up to 40 Non — migrants Teleosts, crustaceans, m depth cephalopods 16. Loliginidae Pelagic, up to 170 Non — migrants Teleosts, zooplankton; m depth may exhibit cannibalism 17. Diogenidae Demersal, coastal Non — migrants Detritus, degrading waters plant and animal matter 18. Astropectinidae Demersal, coastal Non — migrants Juvenile molluscs and waters crustaceans 19. Nemipteridae Demersal, up to 80 Non — migrants Teleosts, wide variety of m depth benthic crustaceans 20. Synodontidae Demersal, up to 60 Migrants Wide variety of teleosts, m depth, some (amphidromous) crustaceans and squids species up to 400 m Table 3.12. Ecological functional guilds of the major fish families observed during the present study

Sr. Family Ecological functional guilds No. Habitat use Vertical Substratum Feeding distribution preference habits 1. Clupeidae Temporary Pelagic — Opportunistic (Marine stragglers, amphidromous 2. Engraillidae Temporary Bentho Mixed Opportunistic (anadromous, — pelagic amphidromous, oceanodromous) 3. Sciaenidae Temporary Bentho Mixed Opportunistic (amphidromous) — pelagic 4. Trichiuridae Temporary Bentho Soft substrate Opportunistic (amphidromous) — pelagic (mud) 5. Portunidae Permanent Demersal Mixed Opportunistic (non — migrants) 6. Tetraodontidae Temporary Demersal Rocky Opportunistic (varied) 7. Lactariidae Permanent Pelagic — Opportunistic (non — migrants) 8. Leiognathidae Temporary Demersal Mixed Opportunistic (amphidromous) 9. Squillidae Permanent Benthic Mixed Zoobenthivore (non — migrants) 10. Penaeidae Permanent Demersal Mixed Opportunistic (non — migrants) 11. Cynoglossidae Permanent Demersal Mixed Opportunistic (non — migrants) 12. Muraenesocidae Temporary Demersal Soft (mud, Opportunistic (oceanodromous) near reefs) 13. Haemulidae Temporary Demersal Mixed (near Opportunistic (amphidromous) reefs) 14. Ariidae Temporary Demersal Soft (mud) Opportunistic (varied) 15. Sepiidae Permanent Demersal Mixed Opportunistic (non — migrants) 16. Loliginidae Permanent Pelagic — Opportunistic (non — migrants) 17. Diogenidae Permanent Demersal Mixed Opportunistic (non — migrants) 18. Astropectinidae Permanent Demersal Mixed Zoobenthivore (non — migrants) 19. Nemipteridae Permanent Demersal Soft Opportunistic (non — migrants) 20. Synodontidae Temporary Demersal Soft Opportunistic (amphidromous)

co oco I I I I I I La On I oti kloo sqt Po o oo Moo Co ro 19 Moo P MVO, MA IMO To ropo Woo imoboo011oo Clop* Moo gro Mao ScIao 11000 TrIcl tr r Moo Porto i Moo TO rraoslo 11Kta0 So p IMoo Loot) rItfoo Cye og boo Moe DOTI I Moo Lough Moo MI roe 108o0100o Ifoom 1 9100 As Ito p o Aritio0 %TOM Int Mon Sling lo Moo Oostopodo N Moo Ocbricl Moo Dooyoltioo I- II ppo 01100 Po VI o M Moo CI Iroco It ido° lootract o In Moo Apia 1100 Sp tyroe I KM re grn oco ro1110o ArCInee Co rcl o rl Il Moo DO rpp Moo Ilyrt tOp I IMO* Stroll Moo Nom lecilltlan Soh Moo MoIldoe Cob nue Pt:NO:leo Pr011gooto Moo Co Mort! Hem IA rktoo odo o Woo Mats Woo Co I'M Moo CO 110 own throe So rro Mao Scorn b 'Moe ✓~pogo l hoe Pa rallolilyKlan 1.1111orrndloY011Os Tilton' It Moo tt l gtiKtae Po mole rtIoe

Fig. 3.19a. Dendrogram indicating similarity among fish families observed during the entire study Stress: 0.18 Hydrophilda e DorIppidae Maybe swop

HIppotytidae HemIs cY111100nemiclagfromateldae Gobiidae Sfftagtntdae Alpheidae Sole' ekangidae TriacantRiVidae Cnidaria Ambasstdae Pristigastertdae SiggrIetchthAdae BrMagraine 11:1-34-1; ae atidae 0+1;,11.:1 ent • ae Arddae Lsi Unidentified juveniles Carcharhinidae Penaeldae Serrantdae Synodonfidae aetirtrartittidVailltileg P arKP e Apogoniclae EttiMERBITIPA

Fig. 3.19b. MDS plot indicating similarity among fish families observed during the entire study CD 2 I 1 I I P bt; ;leo Po leo Mao to ttglati kIDO Am Dm Mao MO taO oeSoollon Hoornt Moo D Ono t kto0 Arikfao lo I Hp Moo p Oleo cytog boa mu Illtta0 Loot ;Moo MOO t Mao TrIol I; Moo Pottle Mae Chpo Mao Ewa; 11100 To traodo; *leo Astropo an boo PrIstignot rklao Paratklifyktoo tippoWniao aostropoclo Mal Mao Tana It; Mao XOtti boo Octpod boo 00%10143o C t Iroco ;trine itpogo; boo Apia kiwi rx)iyron t boo Comer; II Idoo B rogmaao mrttloo Pohroom lope RO tro Moo Co Ilboym boo mob Woo m 011 flIOO 9lt oclotbaoo So ;rat MO Dom b Oleo Arcklao Utkle;tlDedliYai llae Co rolg Icloo Strom at Mae riaticepkottfon $110glo boo Do ;pp Moo Kid ;opt Moo Ho m inyllIctao Solo Mao To repo, Woo Stgai boo Oob Mao CI! MOM

Fig. 3.20a. Dendrogram indicating similarity among fish families observed during the pre — monsoon season Stress: 0.16

600AltadeVe Spnyraentdae AIrtheldae Hippottdae poneinfavalichthytdae Sillagtnidae Mafldae Gerreidae Stromateidae Catlionymidae Platycephatidae.sigantirc rarrettscyMdae Arcfdae MatuUdae ttda gafae igNACcidae, Arnt ,,arangidae Untdentilied juveniles Serrantdae arBI16171atae grow ing ae Scomtrndae T 08ae Dortppidae LettigirintSe Hydrophlidae Penaefdae Apogonidae Cntriaria

Plotosidae

Chfrocentrtdae

Fig. 3.20b. MDS plot indicating similarity among fish families observed during the pre — monsoon season investigate and examine associations during the pre — monsoon season. Cluster II comprised the two most abundant families namely Penaeidae and Leiognathidae (78

% similarity). Cluster III comprised Muraenesocidae and Haemulidae (82 % similarity), and these groups clustered with Ambassidae at 62 % similarity. Cluster IV comprised seasonal associations of fifteen families, among which one sub — cluster namely (a) Diogenidae and Ariidae showed a similarity of > 80 % similarity, and the other three sub — clusters namely (b) Sepiidae, Cynoglossidae and Loliginidae (c)

Squillidae, Lactariidae, Portunidae, Tetraodontidae, Clupeidae, Engraulidae,

Sciaenidae and Trichiuridae, and (d) Astropectinidae and Pristigasteridae showed a similarity of < 80 %. Cluster XI comprised Nemipteridae and Synodontidae (96 % similarity), and these groups clustered with Matutidae at 55 % similarity.

3.3.4c. Family level: post — monsoon season

The cluster analysis of numerical data of fish families for post — monsoon months collected during the present study revealed altogether nine clusters with > 50

% similarity (Fig. 3.21a,b). Among these, only four clusters were selected to investigate and examine associations during the post — monsoon season. Cluster V comprised two sub — clusters namely (a) Ariidae and Leiognathidae with 80 % similarity, and (b) Matutidae and Haemulidae with rz, 75 % similarity. Cluster VII comprised Penaeidae and Squillidae with 85 % similarity. Cluster VIII comprised two sub — clusters namely (a) Tetraodontidae, Muraenesocidae and Synodontidae with

78 % similarity, and (b) Portunidae, Lactariidae, Trichiuridae, Engraulidae,

Cynoglossidae, Clupeidae and Sciaenidae with `;=-1 75 % similarity. Cluster IX comprised Nemipteridae, Terapontidae and Loliginidae with 75 % similarity.

42 co °ts 0 H Iwo hesklaso r rdaatr rKlau ctIclarai Natlaa Co ral art Is Mae Oge t Mae Se tra t Klan Se pills° P laVce pia Mae Sob Mao On tro Klan Scorn I) rIclao Pa ra 101 tlylclaa UlklaltrACdlivatllue /catalpa CtIt Mao MsglItIsse Stia I Klan /Vogl, 1 Klan Pomplo tidos Ca lib 'yin Klan Trlaca l tl Mae Arlklea La Insets' Mao Matilda° Moans I Mae AmPalaslloo CaralgKloo 9q 1 Illklae Pa tan Man To trauclo I Oleo Ms ran IasoCklan SD oar, snarl° Po rim i Mae Loots Slklao TrIcl Is Mae Betas Mae Cyi ag los$ Mae Cli pa Klan Saila l Mae Ne at pis Mac Long's Klan re regain:la°

Fig. 3.21a. Dendrogram indicating similarity among fish families observed during the post — monsoon season A Stress: 0.13 CarctmetAlae

Ambassidae Carangtdae

Nemipterldae Off Mae Terapontidae les Gerreidae Lott • • • , j411/14111:ki. MatutidaglatYcetabletdae REM Soletdae Letognathidae Adidae

PartildffirinOtidae

Fig. 3.21b. MDS plot indicating similarity among fish families observed during the post — monsoon season

4- 3.3.4d. Species level

The cluster analysis of numerical data of the most frequently occurring species

(N = 38) collected during the present study revealed altogether nine clusters with > 50

% similarity (Fig. 3.22a,b). Among these, only five clusters were selected to investigate and examine associations among the dominant species collected during the present study. Cluster II comprised S. tumbil and N. japonicus with 80 % similarity.

Cluster III comprised the ten most abundant coastal residents and migrants those categorized into three sub — clusters namely (a) M. affinis and P. stylifera with 85 % similarity, (b) M. nepa and M. dobsoni with 78 % similarity, and (c) P. bindus, 0. tardoore, T. dussumieri, L. spadiceus, L. lactarius and 7'. lepturus with 70 % similarity. Cluster W comprised demersal coastal residents namely P. pelagicus, P. merguiensis and D. miles with 62 % similarity. Cluster V comprised loosely associated coastal residents and migrant species those categorized into two main sub — clusters namely (a) S. commersonnii, C. macrostomus, C. lucifera, S. longiceps, T purava, A. maculatus, U. duvauceli, S. inermis and Johnius sp. with 62 % similarity, and (b) P. indicus, P. sanguinolentus, S. insidiator, A. indicus, M cinereus and P. maculatus with 55 % similarity. Cluster IX comprised amphidromous species namely P. ditchela, E. splendens and 0. ruber with 61 % similarity.

3.3.4e. Species level: pre — monsoon season

The cluster analysis of numerical data of the most frequently occurring species collected during the present study for the pre — monsoon months revealed altogether eight clusters with > 50 % similarity (Fig. 3.23a,b). Among these, only five clusters were selected to investigate and examine associations during the pre — monsoon.

Cluster II comprised migrant predatory fishes namely S. tumbil, N. japonicus with

43 -a 0 0 N) 0 0 t 0 0 I 1 I 1 I I C. vadorum N. Japonicus S. tumbil M. affinis P. stylifera M. nepa M. dobsoni P. blndus O. tardoore T. dussurnierl L. spadiceus L. lactarlus T. lepturus P. pelagicus P. merguiensis D. miles S. commersonnli C. macrostomus C, lucifera S. longicepa T. purava A. maculatus U. duveuceli S. inermls Johnius sp. P. indlcus P. sangulnolentus S. Insidlator A. Indlcus M.cinereus P. maculatus C, feriata N. nasus T. setlrostris A, gymnocephalus P. ditchele E. eplendens 0 ruber

Fig. 3.22a. Dendrogram indicating similarity among commonly occurring species observed during the entire study Stress: 0.17

P. "teen

IL Vents bobsonl

P. tendus M.neMa

P. clItettela L spadfceus etaftnerftfolus N. names T. PUTAI"4:42114712. mat44514113"" C. vadorum

A. Inreens & mseugkintsuss

T. seerostes S. longleeps D. megiubey

P.nrdcurdnereus E sp1endens C. luctrera P. mergulensts s.turellitleocessus

P. petagteus

A. gyrnnoceptatus

C. fetish

Fig. 3.22b. MDS plot indicating similarity among commonly occurring species observed during the entire study 8 0 C. vadorum N. jeponlcus S. tumbil M. affInIs P. stylltere M. dobsoni P. bindus nepe L. lacterius T. lepturus T. dummied L, epediceus S. commereonnii C. macrostomus P. peleglcus P. merguiensie D, miles C. lucifera S. longiceps C. ferlate N. nasus T. setirostrls U. duvaucell P. sanguinolentus S. inermis Johnius ep, P. Indlcus S. insidiator P. ditchela A Indlcus 0. tardoore T. purava A. maculetus M. cinereus P. maculatue E. splendens A, gymnocephalus o ruber

Fig. 3.23a. Dendrogram indicating similarity among commonly occurring species observed during the pre — monsoon season Mass: 0.16

0.1edala

A. 7/mm6666216s E. 6 plendens T. sylkostis

P. mergutensla P. 1/111121666

nasus 0 tuber C. ludtera M. clnereus O. mites S. longtime P. maadatus

A Indian To.W1916atus S. insidlatat P. Incicus P. ditchela U. diratortentosnitus Job. rt 54ingls C. macrostonts tIARDNit L spacficeus

S. cornmersonnll M. neva P. btmlus C. vadorum

14.40trds AI. 6606661 P. stlitera

Fig. 3.23b. MDS plot indicating similarity among commonly occurring species observed during the pre — monsoon season

'1r" 96 % similarity. Cluster III comprised the most abundant coastal residents and migrant species those categorized into three sub — clusters namely (a) M affinis and

P. stylifera with 88 % similarity, (b) M. dobsoni and P. bindus with 80 % similarity, and (c) M nepa, L. lactarius, T. lepturus, T dussumieri and L. spadiceus with 78 % similarity. Cluster N comprised coastal residents and migrant species those categorized into two sub — clusters namely (a) S. commersonni and C. macrostomus with 75 % similarity, and (b) P. pelagicus, P. mergeuiensis, D. miles,

C. lucifera and S. longiceps with 60 % similarity. Cluster VII comprised common coastal residents and migrants those categorized into three sub — clusters namely (a)

U. duvauceli, P. sanguinolentus, S. inermis and Johnius sp. with 75 % similarity, and (b) P. ditchela, A. indicus, 0. tardoore, T purava and A. maculatus with 65 % similarity. Cluster VIII comprised migrant species those categorized into two sub — clusters namely (a) M. cinereus and P. maculatus with 65 % similarity, and (b) E. splendens, A. gymnocephalus and 0. ruber with 65 % similarity.

3.3.4f Species level: post — monsoon season

The cluster analysis of numerical data of the most frequently occurring species collected during the present study for the post — monsoon months revealed altogether nine clusters with > 50 % similarity. Among these, only seven clusters were selected to investigate and examine associations during the post - monsoon (Fig. 3.24a,b).

Cluster I comprised M. nepa and M dobsoni with r-z-: 80 % similarity. Cluster II comprised two sub — clusters namely (a) coastal residents such as U. duvauceli, C. vadorum and N. japonicus with > 75 % similarity, and (b) migrants such as S. cornmersonnii, M cinereus, S. tumbil, P. maculatus and L. spadiceus with > 65 % similarity. Cluster III comprised mixture of coastal residents and migrant species

44

M. nape M. dobeoni U, duvoucoll O. vedomm N. leponloue S. commorsonnil M. olnorous S. tumbil P. moult:due opedlcous C. luclfore Johnlue ep. C. meoroetomus rubor S. longloope L. lectedue T. lopturue P. sengulnolontus Inoldiotor P. bIndue A meoulotus eplondene P, mornulenslo A Indlous O. tordooro P. potegloue S. Inormlo duoeumtort P. Indlcus D. miles A. gymnocapholue O. fedete P. dttohole P. etyiitore M. effinle T. pureNm T. sotlrostrie

Fig. 3.24a. Dendrogram indicating similarity among commonly occurring species observed during the post — monsoon season

Stress: 0.1T

ttril3Mie"sis

P. trinclussatardo.remiles

E s plendens P. Indian A. macutatus

P. petagicus T. setirostris T. adtennirti a insidiator S. commersonnti purava SIttninftreuV. macutatus

C. feriata P. sangutnotentpatPs sp. laPanirastorum M. affrnis duvaucen P. Oct" eta C. L. snaffle-sus

C. luctiera P. styfifera M. dobsoni gymnocephatus M. nepa

Fig. 3.24b. MDS plot indicating similarity among commonly occurring species observed during the post — monsoon season those categorized into three sub — clusters namely (a) C. lucifera, (b) Johnius sp. and

C. macrostomus with 90 % similarity, and (c) 0. ruber, S. longiceps, L. lactarius and T. lepturus with rz: 80 % similarity. Cluster IV comprised mostly migrant species those categorized into two sub — clusters namely (a) P. sanguinolentus and S. insidiator with 84 % similarity, and (b) P. bindus, A. maculatus and E. splendens with rz 77 % similarity. Cluster V comprised P. merguiensis and A. indicus with 94

% similarity. Cluster VI comprised three sub — clusters namely (a) 0. tardoore, (b) P. pelagicus, S. inermis and 7'. dussumieri with z' 92 % similarity, and (c) P. indicus and D. miles with 84 % similarity. Cluster IX comprised two sub — clusters namely (a)

P. stylifera, and (b) M affinis, T purava and T. setirostris with 78 % similarity.

3.3.5. Catch rates of the study area

The catch rate (kg.hf l) of the study area, particularly the nearshore waters up to 25 m depth was estimated to be 99.47 ± 72.51 kg.hf l. Among the fifteen fish groups, the highest catch rates (30.23 ± 28.75 kg.hf l; Table 3.13) were recorded for the miscellaneous group (crabs, non — edible fin fish, molluscs and other invertebrates), however catch rate of prawns was comparatively lower (21.64 ± 21.47 kg.hfl; Table 3.13). Other important commercial groups such as clupeoids and leiognathids showed lower catch rates (11.51 ± 17.31 and 8.03 ± 22.54 kg.hf 1, respectively). Seasonal trends in fish catches revealed higher catch rates during the post — monsoon (105.23 ± 40.72 kg.hf 1) than during the pre — monsoon (97.07 ± 82.49 kg.hfl). Among the different groups, catch rates of prawns, leiognathids, sciaenids, cephalopods, carangids, pomfrets and elasmobranchs were comparatively higher during the pre — monsoon season, whereas those of miscellaneous groups such

45 Table 3.13. Catch rates of major fish groups during the present study

Sr. Demersal Pre - monsoon Post - monsoon Overall study No. groups (kg. he') % (kg. hr') % (kg. hr') % composition composition composition 1. Stomatopods 9.80 ± 10.09 23.57 ± 22.40 13.85 ± 13.92 14.86 25.38 19.43 2. Prawns 26.06 ± 26.85 11.01 ± 10.46 21.64 ± 21.75 23.93 6.35 21.47 3. Elasmobranchs 0.41 ± 0.43 0.27 ± 0.26 0.37 ± 0.37 1.35 0.86 1.22 _ 4. Clupeoids 10.59 ± 10.91 13.73 ± 13.04 11.51 ± 11.57 11.84 26.52 17.31 5. Sciacnids 4.58 ± 4.72 2.15 ± 2.04 3.87 ± 3.89 13.54 2.10 11.45 6. Lactariids 0.96 ± 0.99 0.99 ± 0.94 0.97 ± 0.98 1.19 1.22 1.20 7. Catfishes 0.69 ± 0.71 1.43 ± 1.36 0.91 ± 0.91 0.99 2.76 1.72 8. Leiognathids 10.92 ± 11.25 1.08 ± 1.02 8.03 ± 8.07 26.35 1.46 22.54 9. Ribbon fishes 2.30 ± 2.37 2.35 ± 2.23 2.32 ± 2.33 4.20 1.97 3.67 10. Flatfishes 1.22 ± 1.26 4.03 ± 3.83 2.05 ± 2.06 1.50 4.57 3.03 11. Carangids 0.56 ± 0.58 0.14 ± 0.14 0.44 ± 0.44 2.65 0.44 2.24 12. Pomfrets 0.50 ± 0.51 0.12 ± 0.12 0.39 ± 0.39 1.50 0.40 1.29 13. Perches 0.47 ± 0.49 0.62 ± 0.59 0.52 ± 0.52 1.03 0.85 0.97 14. Cephalopods 2.79 ± 2.87 1.44 ± 1.37 2.39 ± 2.40 4.57 2.54 4.11 15. Miscellaneous 25.21 ± 25.97 42.28 ± 40.18 30.23 ± 30.39 22.96 37.34 28.75 Total 97.07 ± 105.23 ± 99.47 ± 82.49 40.72 72.51 as stomatopods, clupeoids, flatfishes, ribbon fishes, catfishes, perches and lactariids were higher during the post — monsoon season (Table 3.13).

3.4. Discussion

Coastal ecosystems are endowed with high degree of habitat complexity and dynamic nature due to inherent physico — chemical processes, which enable the establishment of diverse micro — niches those support and sustain variable biota at both microsopic and macroscopic levels.

The present study in the coastal waters of Goa, central west coast of India inventoried two hundred and four taxa. Rao and Dorairaj (1968) attempted to provide an estimate of fishery yield off Goa and listed four genera and one species of commercial fishes. Talwar (1973) reported one hundred and sixty eight fin fish species from the littoral waters of Goa based on the examination of fishes collected by

Dr. S.W. Kemp, Dr. A.W. Herre, Mr. P.F. Jones, the Karnatak College, Dharwar and the Zoological Survey of India, at least thirty years prior to their identification. Tilak

(1973) reported fifty one fin fish species from the rivers and estuaries of Goa. Prabhu and Dhawan (1974) reported forty seven commercially important taxa (ten elasmobranchs, twenty eight teleosts, three stomatopods and six prawns) of demersal marine fauna from the 20 and 40 metre depth regions off Goa coast along with catch rates of few important fishes. George (1980) reported seventeen species of penaeid prawns from the Goa coast along with brief taxonomic descriptions. Ansari et al.

(1995, 2003) reported fifty nine fin fish species (two elasmobranchs and fifty seven teleosts) from the Mandovi — Zuari estuarine complex and the adjacent bays along with seasonal variations in their abundance, distribution, diversity and effect of environmental factors. Lobo (2005) reported eight species of sea snakes from the

46 coastal waters of Goa. Much of the above published literature pertaining to demersal species composition and distribution along the coastal waters of Goa suggested that major focus was laid upon commercially exploited species.

The present study in terms of sampling effort and focus on rare species yielded an addition of fifty five new species to the existing inventory of demersal marine species along Goa coast. These observations highlight that the coastal regions form a congenial habitat for a wide array of marine as well as coastal estuarine species

(Kulkarni et al., 2003). The complexity among the coastal habitats offers diverse niches those support wide array of faunal assemblages (Gratwicke and Speight, 2005).

Further, interplay among the ambient physico — chemical (depth, temperature and salinity) and biological factors (productivity and prey availability) determines the formation of resource — specific species assemblages to enable optimum utilization at different trophic levels within an ecosystem (Elliot et al., 2007).

The observations made on the size class of different demersal fauna indicated the dominance of adults and juveniles of the resident crustaceans, fin fish and other fauna emphasizing that the nearshore waters serve as a spawning as well as nursery grounds for variety of euryhaline fauna (Ansari et al., 1995). Day et al. (1981) suggested that tropical and sub — tropical nearshore areas are the primary spawning grounds for many species, whose juveniles migrate to the estuaries. In addition, the occurrence of young recruits of both migrant shoaling planktivorous and solitary piscivorous fishes suggested that the adjacent bays serve as a foraging ground for these species. Blaber et a/. (1995) emphasized the significance of the shallow inshore waters of Albatross Bay as an important foraging zone for piscivores.

The trends in the fish catch data obtained during the present study revealed the dominance of the crustaceans, as these were target species for bottom trawlers. It

47 is worth mentioning that although the bottom trawl is employed mainly to harvest penaeid prawns, their contribution was only 37.89 and 22.31 % to the total fauna by numbers and weight, respectively. These findings are also in concurrence with the global assessment of demersal fish catches using bottom trawls (Watson et al., 2006), which reveals that the contribution of prawns to the bottom trawl catches reduced significantly from 52.65 % (1970's) to 17.86 % (1990's). The overall decline in prawn catches may be attributed to long — term intensive targeted exploitation using trawl nets with small cod end mesh size, thus leading to recruitment over — fishing as reported along the Goan coast (Ansari et al., 2006). The above trends in the catch of penaeid prawns over past few decades probably play an important role in the reduction of potential predators such as sciaenids and cephalopods in the study area as evident from the low catch rates. The dominance of other groups such as fin fish, other crustaceans, molluscs and other fauna (77.69 %) in the trawl catches during the present study emphasizes the non — selective nature of the fishing gear (Kumar and

Deepthi, 2006). Most of the above fauna, except a few commercially important species, are non — targeted species those are caught incidentally (Watson et al., 2006).

Global estimates of trawl by — catch indicated a two — fold increase from 1970's

(46.84 %) to the 1990's (89.30 %), primarily due to the increased use of bottom trawlers (Watson et al., 2006).

The monthly trends in the trawl catch data obtained during the present study revealed that adults of M. dobsoni and M affinis, juveniles of most fin fish and other crustacean species dominated during pre — monsoon, 2006. The pre — monsoon season is generally characterized by higher temperature, salinity conditions in the nearshore and estuarine waters coupled with increased productivity (Krishna Kumari et al.,

2002), which create conducive environment for foraging by juveniles of most marine

48 fishes. Moreover, the solar prawns are known to be perennial breeders in the Goan coastal waters (Achuthankutty and Parulekar, 1986), and thus form the bulk of the trawl catches at this time. The subsequent reduction in faunal numbers during

December, 2006 suggested the probable migration of most marine fauna to deeper waters for spawning (Potter et al., 1990). However, peaks in the numbers of S. longiceps during the above period suggested their recruitment to the coastal waters.

On the other hand, peaks in stomatopod (M nepa) numbers suggest their proliferation in the absence of potential predators (personal observation), whereas large catches of

L. spadiceus adults suggested spawning migration to the nearshore waters (Naik,

1998). The pre — monsoon season of 2007 was characterized by rough weather conditions those impeded trawling operations along the 10 — 25 m depth contours, resulting in low catches. The dominance of certain species (M. dobsoni, A. indicus, P. bindus, L. lactarius and T. lepturus) during the above period may be attributed to their eco — biological processes. The return of calm sea conditions during November, 2007

— January, 2008 resulted in substantially higher trawl catches characterized by the seasonal dominance of M. nepa, L. spadiceus and clupeoid species. Subsequently, variable sea conditions during the February — May, 2008 resulted in erratic effort those influenced the trawl catches. However, the unusually high average numbers and weight (37.16 ± 28.21 x 103 and 128.29 ± 111.20 kg, respectively) during the above period may be attributed to the trapping of large shoals of P. bindus during March,

2008. Further, analysis revealed the dominance of the crustaceans during most sampling months, thereby indicating that they are the major residents of the coastal waters, whereas the occasional dominance of fin fish during the present study is mainly due to trapping of large shoals of recruits of migrant species as envisaged from the present observations. In addition, the insignificant contribution of the other marine

49 fauna to the trawl catches suggested the role of intrinsic biological characters those regulate their population processes.

The two — way ANOVA revealed significant differences in faunal abundance and weight among faunal groups and suggested high degree of variability in the species composition of individual trawls. The significant differences between the sampling months and seasons may be correlated to the temporal variations in the physico — chemical and biological parameters of the coastal waters those might regulate the biological processes of the resident and quasi — resident species. Similar variations in demersal fish catches were reported by several authors (Bapat et al.,

1972; Prabhu and Dhawan, 1974; Radhakrishnan, 1974; Ansari et al., 1995) along the west coast of India.

The analysis of the crustacean catches revealed that the three major penaeid prawns namely M. dobsoni, M. affinis and P. stylifera generally dominated the catches during the present study. All the above species are residents of the nearshore waters of

Goa, whose juveniles are known to migrate into the adjacent estuaries (Achuthankutty and Parulekar, 1986). However, the occurrence of P. stylifera in lesser quantities during February — May, 2007 is attributed to trawl operations carried out in the depth zone of less than 20 m, which is not a suitable habitat for P. sty!jfera owing to estuarine influence (Achuthankutty and Parulekar, 1986). The reduction in penaeid prawn numbers during March, 2008 may be attributed to the prevailing inclement weather conditions (rough sea) those hindered fishing operations. Further investigation into the crustacean by catch species revealed an inverse relation between pen acid prawns and M. nepa throughout the tenure of the present study. However, more detailed studies involving long — term dietary analyses of these groups are required to substantiate this relationship. The portunid crabs comprised juveniles as

50 well as gravid females throughout the study period thus indicating that these are coastal resident species. However, the low numbers of portunid crabs is suggestive of the inability of the demersal trawl to exploit these resources owing to uneven bathymetry of the potential fishing grounds (Wagle and Kunte, 1999). Further analysis of size class revealed that berried female crabs were observed intermittently during the entire study period, and therefore it was difficult to ascertain the exact spawning period of these species. In addition to these, the occurrence of berried females of two estuarine species namely C. goaensis and S. olivacea during the pre — monsoon indicated spawning migration to the nearshore waters; two other species (S. serrata and 7'. crenata) were collected only from the estuarine embayment indicating that these are essentially estuarine residents; the rare occurrence of C. variegata during the pre — monsoon does not merit plausible explanation owing to lack of data.

The non — portunid crabs, inhabitants of sandy or muddy substrates of coastal waters were rarely caught during the present study. The anomurans were observed in insignificant numbers in the trawl by catch during the present study, and among these only D. miles was observed in comparatively larger numbers in the nearshore waters during the pre — monsoon season. Shenoy and Sankolli (1993) reported that D. miles, a common inhabitant of the Indian sub — tidal region breeds along its western coasts during summer, therefore suggesting breeding aggregation in the coastal waters.

Among the other crustacean by catch groups observed during the present study, the non — penaeid prawns and lobsters occurred rarely probably due to their habitat.

The analysis of fin fish component of the trawl catches taken during the present study revealed different temporal trends among the various constituent fish groups owing to species — specific and group — specific eco — biological requirements

(habitat and food availability) and processes (foraging, breeding and migration). The

51 predominance of juveniles of coastal residents (elasmobranchs and false trevallies) and migrants (clupeoids, catfishes, eels, leiognathids, sciaenids and ribbon fishes) throughout the present study suggested that the nearshore and bay — estuarine waters along Goa coast serve as important nursery grounds for these fauna (Ansari et al.,

1995). Published literature (Fennessy, 2000; Beck et al., 2003; Nagelkerken et al.,

2008) emphasized the importance of nearshore and mangrove — vegetated estuarine regions as nursery grounds owing to nutrient — rich turbid waters coupled with complex habitat structures those provide shelter from tides and wave surges, adequate prey for juveniles, and protection from potential predators due to low visibility. On the other hand, seasonal peaks in the occurrence of adults of coastal residents

(elasmobranchs and flatfishes) and migrants (clupeoids, catfishes, threadfin breams, lizard fishes and puffers) observed during the present study, preferably during

December — February suggested spawning migration to the adjacent bay — estuarine waters. Published literature (Radhalcrisluian, 1963; Rao, 1983; Murty et al., 1992;

Khan and Nandakumaran, 1993; Naik, 1998) pertaining to the spawning activity of demersal marine fishes along the Indian coasts revealed that most marine fishes spawn during the post — monsoon season, and few among these either have an extended spawning period or a secondary one during the remaining part of the year.

Further, the capture of mature individuals of these species from the Mandovi — Zuari estuaries and the nearshore waters suggested that these form congenial spawning grounds owing to the presence of submerged rocks (Wagle and Kunte, 1999) and mangrove vegetation (Kulkarni et al., 2003) those presumably provide adequate substratum and protection to fish eggs. Another noteworthy observation during the present study revealed wide array of "other teleosts" species (N = 74) those formed

52 greater than one — thirds of the total number of species recorded, however they contributed meagrely to the fin fish catches probably due to the habitat complexity.

The analysis of the molluscs and the other fauna catches during the present study revealed that they formed a minor portion of the trawl by catch. Among the molluscs, the cephalopods exhibited some consistency with regards to their temporal distribution patterns with seasonal peaks during March — May suggesting spawning activity in the nearshore waters (Silas et al., 1982, 1986; Jereb and Roper, 2005), whereas the gastropods and bivalves were observed rarely due to submerged ecosystems. The echinoderm catches during the present study were mostly restricted to the pre — monsoon months and the higher catches during April, 2006, March and

May, 2007 are probably due to spawning aggregations in the coastal waters, whereas lower numbers during the other periods suggested migration to deeper waters to avoid strong onshore wave surges (Freeman et al., 2001). The occasional peaks in the numbers of cnidaria during December, 2006 and March, 2008) probably coincided with high densities of zooplankton (personal observation), thereby suggesting foraging migration by the zooplanktivore jellyfish, A. aurita (Papathanassiou et al.,

1987). Further, this phenomenon was observed to coincide with peaks of zooplanktivore fishes (Fig. 3.25).

It is apparent from the previous discussion that the demersal marine fish community described in the present study broadly comprised coastal residents and migrants based on the habitat preference and migratory habits of the individual species. The coastal residents comprising mostly demersal invertebrates

(Achuthankutty and Parulekar, 1986) and solitary fin fish species spend their entire life cycle in the nearshore and bay waters (Ansari et al., 1995). These fauna find adequate shelter among the coastal benthic habitats and protection against strong

53 300 • —•—• Planktivorous fishes • —•—• A. aurita

200

41 100

0 Feb Mar Apr Dec Jan Feb Mar Apr May Nov Jan Feb Mar Apr May I 2006 I 2007 I 2008 Sampling month

Fig. 3.25. Month wise trends in the weights of the jellyfish, Aurelia aurita and planktivorous fishes during the present study wave surges, predation and fishing pressure; their larvae are known to use the turbid nearshore shelf waters (Fennessy, 2000) and the adjacent estuaries (Nagelkerken et al., 2008) as nursery areas owing to food availability, reduction in predation pressure

and relatively calm environment (Blaber and Blaber, 1980).

On the other hand, the migrants comprising mostly shoaling fishes visit the nearshore and bay waters for foraging purpose. The coastal waters of Goa are highly productive (13.12 to 14.21 mg C.n1 3.h-1 ; Krishna Kumari et al., 2002) and sustain high zooplankton biomass (Qasim and Sen Gupta, 1981), which may attract large

shoals of zooplanktivore fishes (Rudstam et al., 1992) during the pre — monsoon; their

larvae and juveniles seek shelter in the turbid waters owing to reduction in predation

pressure and availability of adequate prey (Padate et al., 2010a). Moreover, the

presence of juveniles of migrant piscivores in the study area signifies its role as

nursery area. Similarly, Blaber et al. (1995) reported that the nearshore waters of the

Albatross Bay, Australia served as foraging grounds for piscivores and as nursery

areas for fish juveniles.

The present observations revealed that the demersal faunal assemblages of the

study area are similar to those of other Indo — Pacific regions (Blaber et al., 1989)

those characterized by high species richness (2.04 ± 0.66), diversity (1.94 ± 0.44) and

evenness (0.05 ± 0.02) indicating the stability of the ecosystem (Hooper et al., 2005).

Such high diversity indices seem to be supported by geomorphology and bathymetric

profiles (Veerayya, 1972) coupled with intricacies of the coastal habitats such as

patchy coral reefs (Rodrigues et al., 1998) and mangrove vegetation (Kulkarni et al.,

2003) those provide suitable niches to their inhabitants and enable them to carry out

their respective ecosystem functions through processes such as foraging, breeding,

recruitment and migration. Moreover, artificial structures in the coastal waters such as

54 shipwrecks (Padate et al., 2010a) also enhance the habitat complexity of the coastal ecosystems (Walker et al., 2007) as they readily provide substratum in the form of platforms and crevices for fouling organisms, which further attract colonization by wide array of reef fauna (Arena et al., 2007). Secondly, the physico — chemical processes such as tidal flux, water mass circulation and nutrient recycling play a role through regulation of the spatio — temporal variations among the demersal faunal assemblages. The present values of species diversity were comparatively lower than those reported by Ansari et al. (1995) from the adjacent bay — estuarine region, which may be largely attributed to the dominance of few crustaceans (M. nepa, M. dobsoni,

M affinis, P. stylifera, P. sanguinolentus) and fin fish (P. bindus, L. spadiceus, T lepturus, 0. tardoore, T dussumieri) those contributed > 75 % to the total faunal numbers. Blaber et al. (1989) opined that high diversity and species richness were characteristic of tropical Indo — Pacific estuaries. However, a similar generalization on the nearshore coastal waters cannot be made owing to dearth of studies. De Ben et al. (1990) reported substantially low diversity (0.13 — 1.08) from a temperate bay in

Oregon U.S.A. Araujo et al. (2002) reported values of diversity, species richness and evenness (2.06, 3.17 and 0.74, respectively) from tropical bay waters in Brazil, almost comparable with the present values. Hence, it is imperative that faunal diversity in the tropical nearshore waters of Goa, along the west coast of India is considerably high indicating that diversified habitats along this region play a significant role in the health of the coastal waters. However, more detailed long — term studies need to be carried out to confirm these trends and to elucidate the processes responsible for such phenomena.

Temporal variations in the diversity of demersal fauna during 2006 and 2007 revealed a general increase with the progress of the post — monsoon season followed

55 by a peak during February — March and decrease towards the commencement of the southwest monsoon season. Ansari et al. (1995) reported similar trends in demersal fish diversity from the bay regions of Goa. One of the major causes of high diversity during the present study lies in the increased salinity owing to recession of freshwater

(Qasim and Sengupta, 1981) thereby allowing the return of stenohaline marine migrants to the coastal waters (Ansari et al., 1995). Moreover, phytoplankton blooms during the early pre — monsoon (Nair et al., 1980) may attract large swarms of zooplankton and "young — of — the — year" of planktivorous species (Chen et al.,

2009), which in turn attract foraging by juveniles and recruits of wide array of carnivorous and omnivorous fishes. Secondly, this period coincides with spawning season of a large 'number of demersal invertebrates (Silas et al., 1982, 1986; Roper et al., 1984; Achuthankutty and Parulekar, 1986; Shenoy and Sankolli, 1993; Freeman et al., 2001; Jereb and Roper, 2005) and fin fish (Rao, 1983; Whitehead, 1985, Murty et al., 1992; Naik, 1998). The low values during the post — monsoon season may be explained by reproductive migration among fishes (Annigeri, 1963; Whitehead,

1985). Qasim (1973) opined that most fishes generally spawned during the southwest monsoon (June — September) and the post — monsoon (October — January) along the west coast of India. Further, it is known that most tropical marine species migrate to the deeper coastal shelf waters for spawning purpose (Potter et al., 1990). However, the large fluctuations in the diversity indices during pre — monsoon, 2008 (February —

May), could be attributed to the prevailing rough sea weather enforced by largely uneven meteorological conditions (personal observation).

The cluster analysis of the fish numerical data collected during the present study revealed high degree of similarity among the major fish families. Among these,

Penaeidae and Squillidae (both coastal residents) showed 80 % similarity, thereby

56 suggesting grouping due to common habitat requirements and feeding preferences as evident from similar trophic levels, (2.70 and 3.10), respectively. Further, their grouping with Leiognathidae during the entire study (z . 70 % similarity) and the pre — monsoon season 75 % similarity) is suggestive of the occurrence of prawns coinciding with large shoals of leiognathid juveniles during the above period. On the other hand, the Penaeidae — Squillidae association (> 80 % similarity) during the post

— monsoon season may be attributed to the increase in stomatopod population presumably owing to reduction in predation pressure by migrant fishes, which are known to undertake spawning migration to deeper waters (Potter et aL, 1990). The largest cluster observed during the present study comprised the common migrants

(Clupeidae, Engraulidae, Sciaenidae and Trichiuridae) and coastal residents

(Portunidae). Within this cluster, Clupeidae and Engraulidae (Trophic level: 2.20 —

3.60) are pelagic planktivores, whereas Sciaenidae and Trichiuridae are bentho — pelagic omnivores (Trophic level: 4.00 — 4.30) those forage upon planktivorous fishes and benthic invertebrates (George et al., 1968). Such observations suggested that there would be sharing of prey items between Sciaenidae and Trichiuridae, whereas the Engraulidae may form an intermediate link in the trophic hierarchy that connects the planktivores (Clupeidae) and piscivores (Sciaenidae, Trichiuridae). The other cluster comprising the omnivorous Tetraodontidae (Trophic level: 3.40 — 3.50) and the benthivore Portunidae suggested a prey — predator relationship. The above associations were of seasonal duration as during the pre — monsoon season, most faunal groups migrate to the bay and nearshore coastal waters to forage upon the abundant prey items (Ansari et al., 1995). On the other hand, the post — monsoon cluster comprised diverse faunal groups (Portunidae, Lactariidae, Trichiuridae,

Engraulidae, Cynoglossidae, Clupeidae and Sciaenidae) with a wide array of

57 ecological requirements. Most of the above groups, except Cynoglossidae and

Clupeidae, were represented by juveniles of transient species, which navigate to the estuarine nursery areas probably for shelter and foraging purpose. Another cluster (z

80 % similarity) comprising the predatory omnivores namely Nemipteridae and

Synodontidae (Trophic level: 3.80 and 4.40, respectively), suggested grouping due to similar habitat requirements and feeding habits (Froese and Pauly, 2010). However, the above association was restricted to the pre — monsoon season. In addition to the above, the other clusters namely Muraenesocidae — Haemulidae (> 80 % similarity),

Sepiidae — Cynoglossidae (> 80 % similarity) were seasonal in duration, and based on similar trophic levels (Muraenesocidae: 3.99 — 4.07; Haemulidae: 4.04; Sepiidae:

3.60; Cynoglossidae: 3.26 — 3.50; Froese and Pauly, 2010), suggested sharing of prey comprising benthic invertebrates and small teleosts. The remaining clusters comprised faunal groups with similar dietary requirements (Froese and Pauly, 2010; Palomares and Pauly, 2010) and suggested trophic interactions among them. An assessment of gut contents of these fauna would provide better understanding of the above relationships.

Cluster analysis of the most dominant species observed during the present study revealed season — specificity in the nature of their associations. Among these, two penaeid species namely M. affinis and P. stylifera (coastal residents) showed overall 85 % similarity, and the magnitude of their association was relatively greater during the pre — monsoon (z 88 %) owing to the overlap of their breeding season

(Achuthankutty and Parulekar, 1986). However, low similarity values (r-z: 55 %) during the post — monsoon suggested migration by P. stylifera, a marine species, to the deeper waters probably due to its inability to tolerate relatively low saline conditions

(Menon, 1953) probably caused by the delay in recession of the monsoon during

58 December, 2006 — January, 2007. The cluster comprising M nepa and M. dobsoni (>

80 % similarity), both of which inhabit nearshore waters up to 40 m depth (Palomares and Pauly, 2010), and occupy similar trophic levels (3.1 and 2.7, respectively), suggested grouping due to similar ecological requirements. The cluster comprising the most common opportunistic feeders (Froese and Pauly, 2010) namely P. bindus, 0. tardoore, 7'. dussumieri, L. spadiceus, L. lactarius and T lepturus (> 70 %) suggested complex trophic interactions involving fishes of lower (P. bindus and 0. tardoore) and higher trophic levels (T. dussumieri, L. spadiceus and T lepturus). However, more detailed studies involving analyses of gut contents are required to confirm these associations. Another cluster comprised highly predatory omnivorous fishes namely

S. tumbil and N. japonicus 80 % similarity) those inhabit soft mud substrates, and their feeding habits (Froese and Pauly, 2010) suggested that they share similar food resources. However, the seasonal nature of their association (> 90 % similarity during pre — monsoon) may be attributed to the overlapping of their spawning seasons as S. tumbil is a perennial spawner (Tiews et al., 1972), whereas N. japonicus spawns from

September to May (Murty et aL, 1992). Other clusters comprised loosely associated

(z 52 % similarity) coastal residents (C. lucifera, P. sanguinolentus, S. inermis, U. duvauceli, A. indicus and C. macrostomus) and migrants (S. commersonnii, S. longiceps, T. purava, A. maculatus, S. insidiator, M. cinereus and P. maculatus), and the existence of such mixed assemblages is probably due to the euryhaline nature of these species (Ansari et al., 1995), which enables them to navigate from marine waters to the estuaries and back.

The present assessment of the marine fish productivity of the coastal waters of

Goa revealed catch rates to be 99.47 ± 72.51 kg.hf l . The comparison of the present values with published literature from Goa coast (151 — 200 kg.hf l ; Prabhu and

59 Dhawan, 1974) indicated substantial decline of 51.27 % in the fishery potential of the above region in the last three decades. Further, the present values are much lower than those reported from the neighbouring Karwar coast (192 kg.hf l; Bapat et al., 1972) and those estimated for the entire west coast of India (100 — 800 kg.hr -1 ; Lowe —

McConnell, 1987). The overall decline in the catch rates of demersal fish off Goa may be attributed to a significant increase in the fishing effort by mechanized fishing vessels. The latest figures in terms of mechanized fishing vessels stood at 1157

(Department of Fisheries, Government of Goa, 2007). The above increase in the fishing effort stemmed from the rising demand for commercial species such as M dobsoni from the lucrative international market (Sonak et al., 2006), which presumably led to over — stepping of the maximum sustainable yield (MSY) for the

Goan waters (Ansari et al., 2006).

Further computation of catch rates of important demersal fish during the present study revealed lower values (Table 3.14) as compared to those reported by

Prabhu and Dhawan (1974) along with significant reduction of 28.72 — 74.38 % and

35.85 — 94.15 % in the catches of top carnivores (elasmobranchs, carangids and cephalopods) and omnivores (catfishes, pomfrets, sciaenids and lactariids), respectively. On the other hand, a phenomenal increase was observed in the catches of miscellaneous demersal fauna (1873.27 %), perches (478.34 %) and leiognathids

(249.38 %). The reduction in catches of carnivores and omnivores coupled with increasing catches of planktivores and detritivores during the present study indicated gradual replacement of higher trophic level species with those by lower trophic levels

(Essington et al., 2006). Such long — term shifts in the catch composition are suggestive of concerted efforts to target commercially important species, thereby leading to reduction in the stocks of dominant commercial species, and probable

60 Table 3.14. Catch percentage of major fish groups off Calangute - comparison with Prabhu and Dhawan (1974)

Sr. No. Fish groups Catch percentage (of the total) % change Prabhu and Present study Dhawan (1974) 1. Stomatopods 11.62 13.92 +19.80 2. Prawns 25.88 21.75 - 15.96 3. Elasmobranchs 1.46 0.37 - 74.38 4. Clupeoids 14.37 11.57 - 19.45 5. Sciaenids 13.15 3.89 - 70.42 6. Lactariids 1.52 0.98 - 35.85 7. Catfishes 15.56 0.91 - 94.15 8. Leiognathids 2.31 8.07 +249.38 9. Ribbon fishes 2.95 2.33 - 21.04 10. Flatfishes 1.89 2.06 +8.97 11. Carangids 1.02 0.44 - 56.59 12. Pomfrets 3.28 0.39 - 88.08 13. Perches 0.09 0.52 +478.34 14. Cephalopods 3.37 2.40 - 28.72 15. Miscellaneous 1.54 30.39 +1873.27 replacement by resilient species of low commercial value. Apart from the direct effects of fishing, other factors such as degradation in water quality and habitat alteration may disturb the demersal fish community structure and modify its functioning (Blaber et al., 2000), leading to change in the overall trophic levels of the coastal waters. Nagai (2003) reported very high densities of jellyfishes during the

1990's in the Seto Inland Sea, Japan owing to high eutrophication levels, which led to overall reduction in the trophic levels of the demersal fish community.

3.5. Conclusion

The present observations along the nearshore and bay — estuarine waters of

Goa revealed altogether two hundred and four taxa belonging to four major faunal groups. Among these fifty five were new records to this region, including one new species to science and four others first records to the west coast of India. The quantitative analysis of the trawl catches revealed the dominance of crustaceans, however there was reduction in the penaeid prawns catches. High species diversity values recorded during the present study indicated the stability of the ecosystem. A comparison of the present fish catch rate (99.47 ± 72.51 kg.hf l) with earlier published literature indicated a sharp decline (51.27 %) during the past three decades, attributed to significant rise in the fishing effort. Further, reduction in catches of higher carnivores coupled with phenomenal increase in lower trophic level catches reflected gradual replacement by planktivores.

61 Chapter 4. Description of Charybdis (Charybdis) goaensis, new species 4.1. Introduction

Swimming crabs (Family Portunidae Rafinesque — Schmaltz, 1815) are crustaceans those inhabit sub — tidal, estuarine and offshore waters, and are widely distributed across the Indo — West Pacific region (Stephenson, 1972). Family

Portunidae is a large taxonomic group comprising four hundred and four species belonging to seven sub — families (Caphyrinae, Carcininae, Carupinae,

Podophthalminae, Polybiinae, Portuninae and Thalamitinae) and forty genera (Ng et al., 2008). Among these, Thalamitinae Paulson, 1875 constitutes the largest sub — family comprising one hundred and fifty three species belonging to four genera namely Charybdis, Gonioinfradens, Thalamita, and Thalamitoides. Charybdis De

Haan, 1833, is the second largest genus within the Thalamitinae with sixty species as compared to Thalamita Latreille, 1829 (eighty nine species) (Ng et al., 2008) distributed among four sub — genera (Charybdis De Haan, 1833; Goniohellenus

Alcock, 1899; Gonioneptunus Ortmann, 1893; Goniosupradens Leene, 1938).

4.2. Literature review

For the past two centuries, naturalists have contributed immensely to the taxonomy of the genus Charybdis (De Haan, 1833) through extensive study of brachyuran fauna. Most species were identified based on fauna collected from deep — sea and coastal expeditions throughout the Indo — West Pacific region (Linnaeus,

1758; Fabricius, 1787, 1798; Herbst, 1789, 1801; de Haan, 1833, 1835; Hombron and

Jacquinot, 1846; Dana, 1852; A. Milne Edwards, 1861, 1869; Miers, 1879, 1884,

1886; Alcock, 1899a; Nobili, 1906; Parisi, 1916; Rathbun, 1923a, 1923b, 1924; de

Man, 1925; Gordon, 1931; Shen, 1932, 1934, 1937; Chopra, 1935; Leene, 1937,

1938; Ward, 1941; Edmondson, 1954; Stephenson et al., 1957; Rees and Stephenson,

62 1966; Stephenson and Rees, 1967b, 1968; Zarenkov, 1970; Sakai, 1976; Crosnier,

1984). In addition, some species were also identified from museum collections (Leene and Buitendijk, 1949; Stephenson, 1972). Among these, Leene (1938) provided detailed descriptions of thirty three species and seven varieties including three new species namely C. amboinensis, C. rathbuni and C. salehensis. Further, she re — organized the genus Charybdis through the introduction of two new sub — genera namely Goniosupradens and Gonioinfradens to the existing three sub — genera

(Charybdis, Goniohellenus and Gonioneptunus). Stephenson (1972) provided taxonomic key to swimming crabs of the genus Charybdis in view of its extensive nature encompassing forty nine species from the Indo — West Pacific region.

Recent studies pertaining to Charybdis crabs from the Indian Ocean region were initiated by the "Discovery" expedition (Stephenson and Rees, 1967a) which revealed one previously known species namely C. smithii. Crosnier (1984) identified

C. seychellensis from the Mascarene Islands. Subsequently, six expeditions in the western Indian Ocean region ("RN Akademic Petrovsky", "RN Odissey", "R/V

Akademic Kurchatov", "R/V Meteor", "R/V Professor Mesyatsev", "RN Vitiaz") reported nine species in which two (C. crosnieri and C. meteor), were new to science

(Neumann and Spiridonov, 1999; Spiridonov and Tiirkay, 2001). Spiridonov (1999) reported five species from the Rumphius Biohistorical Expedition to Ambon

(Indonesia). The "Anambas" expedition in the eastern Indian Ocean region revealed one previously known species, C. truncata (Yeo et al., 2004).

In addition to the above expeditions, several local surveys were also undertaken in the above region. Wee and Ng (1995) reported nineteen species from the coastal waters off Malaysia and Singapore. They provided detailed morphological descriptions of these species based on the examination of specimens deposited at the

63 local natural history museums (Wee and Ng, 1995). Apel and Spiridonov (1998) reported seventeen Charybdis species from the Arabian Gulf, the Gulf of Oman, southern Oman and Pakistan based on investigation of fresh as well as museum collections. Vannini and Innocenti (2000) reported six species from Somalia and neighbouring coasts.

A review of published literature revealed that out of forty three extant species of the sub — genus Charybdis, thirty five species inhabit estuarine and nearshore shallow waters (Alcock, 1899a; Leene, 1938; Ward, 1941; Wee and Ng, 1995), three inhabit deep — sea as well as shallow coastal environments (Stephenson and Rees,

1968; Spiridonov and Tiirkay, 2001) and five species inhabit deep — sea environments

(Moosa, 1995; Spiridonov and Tiirkay, 2001).

In addition to the extant species of the genus Charybdis, published reports (Hu and Tao 1979, 1985, 1996, 2000; Morris and Collins, 1991) described five extinct species (C. kilmeri Hu, 1984; C. monsoonis Hu & Tao, 1985; C. feriatus amenia Hu

& Tao, 2000; C. gigantica Hu & Tao, 2000; C. feriata bruneiensis Morris & Collins,

1991) from east and southeast Asia based on diagrammatic reconstruction of fossilized remnants of crabs. However, these species were not compared with the extant species in view of the ambiguity in their description.

The present chapter describes a new species, Charybdis (Charybdis) goaensis from the estuarine and nearshore waters of Goa, west coast of India (Padate et al.,

2010b). Further, the new species is compared with all the existing congeners and an updated identification key incorporating forty three species of sub — genus Charybdis is provided (Padate et al., 2010b).

Terminology used in the morphological description of the new species follows

Wee and Ng (1995). In addition, terminology describing thoracic sternum and

64 abdominal cavity follows Ng (2000). The following abbreviations are used: CL — carapace length; CW — extreme width of carapace; LC — length of left cheliped; RC — length of right cheliped; OD — orbital diameter; IS — inter — orbital space / Frontal width; FW — Fronto — orbital width; G1— male gonopod / First pleopod.

Altogether, seven morphometric parameters (CW, CL, OD, IS, FW, LC and

RC) were measured using vernier callipers with an accuracy of 0.01 mm. The three ratios namely CW / CL; OD / IS and IS / CW were derived. In the case of chelipeds, the measurement included segments namely dactylus, propodus, carpus and merus.

The gonopods were removed from the paratype male for Scanning Electron

Microscope (SEM) photography to ascertain the identity and distinctiveness of the species. In addition, Camera lucida diagrams of the ventral surface of male and female specimens were drawn to illustrate the thoracic sternites and the abdominal cavity of male and female specimens, respectively. The type specimens of the new species are stored in 5 % buffered formalin (buffered with hexamethylene tetramine to prevent fragmenting of appendages) and are deposited at the Marine Biology laboratory, Department of Marine Sciences, Goa University.

4.3. Taxonomy

4.3.1. Family Portunidae Rafinesque — Schmaltz, 1815

The Family Portunidae is characterized by flat sub — hexagonal or transversely oval carapace that is usually broader than long, its frontal margin broad, horizontal, lobulate or dentate; presence of characteristic "portunid lobe" on endopodite of first maxilliped; distal two joints of last pair of legs usually flatly expanded; abdominal segments 3 — 5 generally fused in males; sexual opening coxal in males, sternal in females (Apel and Spiridonov, 1998).

65 This group is further divided into seven sub — families namely Caphyrinae Paul'son, 1875; Carcininae MacLeay, 1838; Carupinae Paul'son, 1875; Podophthalminae Dana, 1851; Polybiinae Ortmann, 1893; Portuninae Rafinesque —

Schmaltz, 1815; Thalamitinae Paul'son, 1875 (Apel and Spiridonov, 1998; Ng et al., 2008).

4.3.2. Subfamily Thalamitinae Paul'son, 1875

The sub — family Thalamitinae is characterized by sub — hexagonal or sub — trapezoidal carapace that is markedly broader than long, its antero — lateral margin divided into four to seven teeth; basal antennal segment transversely broadened, usually filling orbital hiatus, with ridges, granules or spines; antennal peduncle and flagellum usually excluded from orbit; chelipeds longer than ambulatory legs, bearing a set of spines on merus, carpus and manus; last pair of legs with paddle — shaped propodi and dactyli; 01 with sub — terminal spines (Apel and Spiridonov, 1998).

This group comprises four genera namely Charybdis De Haan, 1833;

Goniohellenus Leene, 1938; Thalamita Latreille, 1829; Thalamitoides A. Milne

Edwards, 1869 (Ng et al., 2008).

4.3.3. Genus Charybdis De Haan, 1833

Charybdis, De Haan 1833, pp. 9 — 10. (type species: Cancer sexdentatus Herbst, 1783

(a junior subjective synonym of Cancer feriatus Linnaeus, 1758))

The genus Charybdis is characterized by the fronto — orbital border being decidedly less than the greatest width of the carapace; antero — lateral borders oblique and arched, cut into six or seven teeth (Sakai, 1976; Wee and Ng, 1995; Fig. 4.1a); G1 tubular, curved or bent in distal part, tip usually elongated or truncated, with spines on

66 Fronto-orbital margin

Distally bent Spines on Six antero-lateral GI lateral margin teeth

Carapace width (a) (b)

Fig. 4.1. Generalized schematic diagram of the genus Charybdis indicating distinguishing morphological characters (a) Dorsal surface of carapace, (b) G1 lateral surfaces and marginal spinules inside opening, along ventral and dorsal margins (Apel and Spiridonov, 1998; Fig. 4.1b).

4.3.4. Charybdis (Charybdis) goaensis sp. nov.

4.3.4a. Material examined

Holotype: male (designated here GUMS — 1), 29 September 2006, CL 12.80 mm, CW

24.32 mm, Zuari estuary, Goa, central west coast of India (between 15°24'41.2"N,

73°48'56.6"E and 15°24'41.9"N, 73°51'11.0"E), 6 — 7 m depth, demersal trawl.

Paratype: Zuari estuary, Goa: one mature female (GUMS — 2), CL 15.10 mm, CW

24.52 mm; one male (GUMS — 3), CL 21.88 mm, CW 34.08 mm; one male (GUMS —

4), CL 14.02 mm, CW 25.10 mm; one immature female (GUMS — 5) CL 11.92 mm,

CW 22.12 mm (location and date of collection same as holotype); nearshore fishing grounds, Goa: one ovigerous female (GUMS — 6) CL 19.40 mm, CW 29.92 mm, nearshore trawl catch, Goa, India, (between 15°29'43.2"N, 73°44'39.7"E and

15°33 '15.2"N, 73°42'41.6"E), 11 March 2007.

4.3.4b. Diagnosis

Carapace evenly tomentose / pilose, with the exception of transverse ridges and swellings. Median frontal teeth bluntly triangular, sub — medians and lateral frontal teeth separated by wide U — shaped notch. Orbits with strong dorsal inclination; major orbital diameter two — fifths to nearly half the length of the inter — orbital space. Basal antennal joint with smooth granular ridge. Antero — lateral teeth six in number, first slightly notched, sixth greater than two times the length of the preceding five and laterally directed; teeth separated by wide notches. Transverse ridges of carapace distinctly granular, two proto — gastric, one meso — gastric, a

67 medially divided meta — gastric, and epibranchials along the level of the last antero — lateral teeth; no transverse ridges beyond the level of the last antero — lateral teeth; prominent tuberosity on cardiac region covered with densely spaced small granules.

Chelipeds subequal in both sexes, rough, unevenly tomentose; merus with two well — developed spines and five to six granules on anterior margin; manus tumid, five —

costate, with five spines on upper surface, two on distal margin immediately posterior to articulation with dactylus possess blunt tips; dactylus smooth, strong and slender

with curved tips, longer than manus. Propodus of natatory leg smooth, merus bears

prominent spine on postero — distal margin. Sixth abdominal segment in male broader

than long, its sides parallel over less than half their lengths and gradually converge

distally. G1 L — shaped, its distal tip short, stout, with a slight bend; membrane

commences shortly behind the tip; inner row of four to five bristles very sparsely

placed commencing distantly from distal tip and terminate before outer row of

brit-Ales. Female gonopore slightly reniform and located on thoracic sternite 6,

posterior to suture 5/6 and corresponds to the bases of second pair of pereiopods.

4.3.4c. Description

Carapace (CW p, = 26.68 ± 4.44 mm; CL = 15.85 ± 3.94 mm; CW / CL =

1.71 ± 0.16) evenly tomentose / pilose, with the exception of transverse ridges and

swellings (Figs. 4.2a, 4.2b). Frontal margin cut into six teeth (excluding inner supra —

orbital angles), median frontal teeth bluntly triangular, projecting beyond broadly

rounded sub — medians, laterals triangular, narrowest, separated from sub — medians

by wide, U — shaped notch (Fig. 4.2c). Orbits (ODµ = 3.05 ± 0.56 mm) with strong

dorsal inclination; major orbital diameter two — fifths to nearly half of inter — orbital

space (ISµ = 6.26 ± 1.52 mm; OD / ISµ = 0.49 ± 0.03). Inter — orbital space almost

68 10mm 10 mm

(i) Swellings on carapace devoid of tomentum (ii) Tomentose carapace

(a) (b)

(I)

(i) Wide U-shaped notch separating sub-median and lateral frontal teeth (ii) Bluntly triangular median frontal teeth

(c)

Fig. 4.2. Charybdis (Charybdis) goaensis sp. nov. (a) Dorsal surface of carapace (Camera lucida diagram), (b) Dorsal surface of carapace (Photograph), (c) Frontal margin of carapace (Camera lucida diagram) one — fourths carapace width (IS / CW µ = 0.23 ± 0.02). Fronto — orbital margin (FW = 12.77 ± 2.85 mm) almost half carapace width; antero — lateral margins oblique and more or less arched. Infra — orbital lobe with row of granules on frontal margin. Antennae fold transversely. Basal antennal joint with smooth granular ridge, its prolongation in contact with front, thus excluding antennal flagellum from orbit. Merus of third maxilliped with outer distal angle produced sideways. Antero — lateral margin of carapace moderately arched, cut into six serrated teeth separated by wide notches (Fig. 4.2d). First tooth slightly notched, second and third sub — equal, blunt pointed with serrated margins, fourth and fifth wider and spiniform, sixth tooth greater than twice the length of the preceding five and laterally directed (Fig. 4.2d). Posterior margin of carapace slightly curved, forms a curve with the postero — lateral margins of carapace. Transverse ridges distinctly granular, two proto — gastric, one meso — gastric, a medially divided meta — gastric, and epibranchials along the level of the last antero — lateral teeth; frontal ridge obsolete. No transverse ridges beyond the level of the last antero — lateral teeth; prominent tuberosity on cardiac region covered with densely spaced small granules (Figs. 4.2a, 4.2b). Surface of transverse ridges and tuberosities more or less devoid of tomentum. Chelipeds subequal in both sexes (LC 29.30 ± 7.36 mm; RC 29.04 ± 8.51 mm), rough, tomentose, a little less than twice the carapace length (Fig. 4.2e). Right cheliped larger in male, left cheliped larger in ovigerous female. Upper surface of merus granular with two well — developed spines and five to six granules on its anterior margin; no spine on posterior margin. Carpus with tuberculated upper surface bears one large spine on its inner angle and three small spines on outer angle. Manus tumid, five costate, with five spines on its upper surface; two blunt spines on distal

69 (i) First antero-lateral tooth slightly notched (ii) Wide notches separate the antero-lateral teeth (iii) Last antero-lateral tooth largest and laterally directed (d)

(I)

c— r CSC r cc--C' z"

1 mm

(i) 2 spines with blunt tips on upper surface of manus (ii) 3 well-developed spines on upper surface of manus (iii)2 spines on anterior margin of mews

(e)

Fig. 4.2. Charybdis (Charybdis) goaensis sp. nov. (d) Antero — lateral margin of carapace (Camera lucida diagram), (e) Dorsal view of right cheliped (Camera lucida diagram) margin immediately posterior to articulation with dactylus; two well — developed spines in the middle of upper surface trailed by coarse granular costae; fifth spine on the posterior margin just anterior to articulation with carpus; outer surface with three smooth costae, the lowermost reaches tip of dactylus. Dactylus smooth, strong, and slender with curved tips, longer than manus. Ambulatory legs long, slender; third pair longest, although, shorter than the chelipeds. Dactyli and propodi of first three pairs of legs covered with densely packed bristles on both anterior and posterior margins, carpus and merus bear bristles only on posterior margin. Posterior margin of propodus and carpus of natatory leg smooth, merus with prominent spine on its postero — distal margin. Thoracic sternites and abdominal cavity in male sternum finely granular (Fig. 4.2f). First three thoracic sternites narrow, sutures 1/2 and 2/3 prominent. Suture 3/4 wide and shallow. Sternite 4 widest, medially divided by a shallow anterior extension of abdominal cavity. Abdominal cavity commences from posterior portion of sternite 4 and covers sternites 5 — 8. Deep groove along abdominal cavity medially divides sternites 5 — 8. Granule of abdominal lock located immediately anterior to suture 5/6. Thoracic sternites in female specimens similar to those in male (Fig. 4.2g). Sixth abdominal segment in males broader than long, its sides parallel over less than half their lengths and then gradually converge distally (Fig. 4.2h); basal margin slightly convex, distal margin concave. Females exhibit dimorphism in abdominal shape; immature females with narrow, triangular abdomen (Fig. 4.2i), wide watch glass — shaped abdomen in ovigerous female (Fig. 4.2j). Female gonopore slightly reniform, located on thoracic sternite 6, posterior to suture 5/6 and corresponding to the bases of second pair of pereiopods (Fig. 4.2g).

70 (V) (I) (r)

(iii)

5mm (iv) 5 mm

(i) Grain of abdominal lock (ii) Thoracic sternites 1-3 (iii) Thoracic sternites 4-8 (iv) Abdominal cavity (v) Female gonopore pair (f) (9)

1--I 1--I 1mm 1mm mm (i) Sixth abdominal segment (h) (i ) (i)

Fig. 4.2. Charybdis (Charybdis) goaensis sp. nov. (f) Ventral surface indicating thoracic stemites and abdominal cavity of male (Camera lucida diagram), (g) Ventral surface indicating thoracic sternites and abdominal cavity of female (Camera lucida diagram), (h) Abdomen of male (Camera lucida diagram), (i) Abdomen of immature female (Camera lucida diagram), (j) Abdomen of mature female (Camera lucida diagram) G1 L — shaped, its distal tip short, stout, with a slight bend (Fig. 4.3a), reaches sixth thoracic sternite; membrane commences shortly behind the tip; outer row of bristles commences close to distal tip and terminates in the neck region; bristles clumped .towards the distal end and sparsely placed towards the neck; inner row of four to five bristles very sparsely placed commencing distantly from distal tip and terminate before outer row of bristles; bristles on abdominal margin randomly and very widely placed (Figs. 4.3b, 4.3c).

4.3.4d. Colour

Carapace light brown over tomentose regions, polished surfaces of granular transverse ridges and regional tuberosities dark brown; pale brown to off — white ventrally. Chelipeds dark brown with whitish tips; other pereiopods light brown.

4.3.4e. Etymology

The species name, Charybdis (Charybdis) goaensis is derived from the type locality, Goa, west coast of India.

4.3.4f Distribution

Charybdis (Charybdis) goaensis is currently known only from the type locality (Zuari estuary) and nearshore fishing grounds of Goa, west coast of India.

4.3.4g. Habitat characteristics

The specimens were collected from 6 — 7 m depth localities from the Zuari estuary, Goa, west coast of India during September (end of the southwest monsoon).

At the time of collection, salinity recorded was in the range of 22 — 32 ppt. The

71 Fig. 4.3. Charybdis (Charybdis) goaensis sp. nov. Scanning Electron Micrographs (SEM) of First left pleopod or Gonopod (G1) (a) Entire Gi, (b) Tip of Gl, (c) Tip of G1 (enlarged view) collection site is in the close vicinity of Mormugao Port Trust, and the waterway is subjected to regular dredging.

4.3.417. Habit

The new species (C. goaensis) was mainly collected from a trawl operation in the estuary. However, capture of an ovigerous female specimen in a nearshore trawl from a depth of 13 — 14 m in March 2007, indicates the probability of catadromous habit of the females of this species for spawning.

4.3.4i. Remarks

Crabs were identified up to the sub — genus level using orthodox taxonomic studies based on morphological characters described by Alcock (1899a), Leene

(1938), Sakai (1976) and Wee and Ng (1995). The distinguishing morphological characteristics of the sub — genus Charybdis are: lobule at external angle of basal antennal joint in contact with front and completely excludes flagellum from orbital hiatus; posterior angles of carapace may be accented or not, but posterior margin forms a curve with postero — lateral margins; four median frontal teeth not dissimilar to lateral frontal teeth; antero — lateral margins cut into six or seven teeth (six large and one small teeth); no spine on posterior margin of cheliped (Leene, 1938; Sakai,

1976).

The distinctiveness of the new species was ascertained based on the comparison with morphological descriptions of its congeners (Fabricius, 1787; Miers,

1884; Alcock, 1899a; Rathbun, 1923a, 1923b, 1924; Shen, 1934; Leene, 1938; Ward,

1941; Leene and Buitendijk, 1949; Edmondson, 1954; Rees and Stephenson, 1966;

Stephenson and Rees, 1967b, 1968; Zarenkov, 1970; Sakai, 1976; Crosnier, 1984;

72 Wee and Ng, 1995; Spiridonov and Tiirkay, 2001). Morphological characters such as number of antero — lateral teeth, presence of transverse ridges on carapace, armature of chelipeds and natatory legs were used as criteria to differentiate the new species

from its congeners. Based on the above criteria, C. goaensis with six antero — lateral

teeth differed with C. heterodon and C. demani with five and seven antero — lateral

teeth, respectively. The new species differed with twelve other species (C. rathbuni,

C. salehensis, C. variegata, C. brevispinosa, C. granulata, C. natator, C.

seychellensis, C. beauforti, C. curtilobus, C. cookei, C. rostrata and C. callianassa)

by the absence of transverse ridges 'posterior to last antero — lateral teeth. Further, C.

goaensis with two spines on anterior margin of merus of cheliped differed with

twenty three other species (C. orientalis, C. hawaiensis, C. feriata, C. padadiana, C.

rosaea, C. annulata, C. hellerii, C. vannamei, C. spinifera, C. acuta, C. incisa, C. javanensis, C. lucifera, C. amboinensis, C. yaldwyni, C. jaubertensis, C. japonica, C.

affinis, C. acutidens, C. rufodactylus, C. sagamiensis, C. riversandersoni, C. crosnieri

and C. holosericus) with three spines, and two species (C. miles and C. meteor) with four to five spines, respectively.

The new species was observed to be morphologically most similar to C.

philippinensis, C. ihlei and C. anisodon based on "two spines on anterior margin of merus of cheliped", and "absence of spines on posterior margin of propodus of

natatory leg". Further, both C. goaensis and C. philippinensis possess "slightly notched first antero — lateral tooth" and "five spines on cheliped manus" and differed from "obtusely triangular first antero — lateral tooth" and "two spines on cheliped

manus" in C. anisodon, and "distinctly bifid first antero — lateral tooth" and "three

spines on cheliped manus" in C. ihlei.

73 The identification of the new species was further confirmed through thorough

comparison of its morphological characteristics with those of the holotype (AMNH

8382) and male paratype (AMNH 7893) specimens of its closest congener, C. philippinensis those obtained from the American Museum of Natural History

(AMNH), New York.

4.3.5. Charybdis (Charybdis) philippinensis Ward, 1941

4.3.5a. Original description by Ward (1941)

"The carapace is broader than long; bare and glossy, granulated under lens. Four

ridges across the carapace placed as in C. vannamei; the first is obsolete and the

others are less developed than in C. vannamei." (In C. vannamei (Ward, 1941), "there

are four granulated ridges on the anterior half of the carapace; the first is placed on the

epigastric region and is broken into two by a broad shallow sulcus which extends back

from the incision between the median pair of frontal teeth; the second ridge is also

divided into two and more widely separated than the epigastric ridges. The third ridge

is entire and concave. The fourth ridge extends from the last tooth of the anterolateral

margin and is broken on each side of the gastric region at the cervical groove.")

"The front is equal to one — fourth the width of the carapace and is cut into six

teeth excluding the inner orbital angles. The sub — median teeth are the broadest,

truncated and sloping toward the median teeth, the lateral pair is narrow obtuse and

sloping inward."

"The anterolateral margins are armed with six spines, including the outer

orbital angle; the spines become larger posteriorly; the last is transversely directed and

twice as long as any of the others. The posterolateral margins are strongly convergent

and outlined by a fine ridge which ends abruptly close to the raised edge of the

74 epimeral wall between the fourth and fifth ambulatory legs. A strong ridge outlines the edge of the epimeral wall and continues unbroken into the posterior margin of the carapace."

"The orbits are small, dorsally directed so that the entire lower margin is visible from a dorsal view when the eyes are retracted; two obsolete fissures in the upper margin, one below the outer angle. The outer angle is formed by a tooth similar in shape to that in C. cruciata (Herbst). The orbital hiatus is filled by the prolongation of the base of the antenna but is extremely short."

"The epistome is well developed though sunken. The buccal orifice is completely closed by the external maxillipeds. The external maxillipeds resemble those of C. vannamei." (In C. vannamei, "the merus of the external maxillipeds, is equal to half the length of the ischium, the outer angle is produced into a narrow auriculate angle. There are two broad sulci on the outer surface with patches of granules and coarse golden hairs near the margins. The palp is large and fringed with coarse golden hairs. The ischium is flat and flossy with a deep longitudinal sulcus parallel to the inner margins.")

"The chelae are subequal. The merus is armed on the anterior margin with two spines and granules, the latter continue onto the upper surface and form a distinct patch becoming reduced in size toward the center. The carpus is armed with four spines, the largest of which is on the inner angle; the upper surface is granulated. The manus is very much inflated proximally and armed above with two spines placed side by side at some distance from the articulation of the dactylus; strong ridges of granules extend posteriorly from these spines. Three broad and low ridges extend longitudinally along the outer surface. The dactylus is equal to the length of the upper border of the manus; both fingers are strongly curved inward toward the front of the

75 carapace, the teeth on both fingers are tricuspidate and interlocking and the tips cross when gripping."

"The ambulatory legs are long and slender; the distal articles are fringed with long hair."

The above description by Ward (1941) was found to be lacking in morphometric and other minute details. In view of the above, the type specimens obtained from AMNH were re — examined and a brief updated description of C. philippinensis along with its morphometry and diagnostic morphological illustrations

(Camera lucida diagrams and photographs) is provided below.

4.3.5b. Updated description of type specimens — C. philippinensis

The updated description is based on comparison of description of Ward (1941) and re — examination of holotype specimen (Catalogue No. AMNH 8382) obtained from the American Museum of Natural History, New York. The holotype specimen obtained was devoid of the right cheliped and legs on right side (Figs. 4.4a — d).

Carapace (CW µ = 35.65 ± 2.56 mm; CLµ = 20.50 ± 1.30 mm; CW / CLµ =

1.74 ± 0.01 mm) naked, devoid of tomentum (Fig. 4.4a). Median frontal teeth rounded, sub — median and lateral frontal teeth separated by narrow V — shaped notch

(Fig. 4.4c). Orbits with strong dorsal inclination; major orbital diameter (ODµ = 3.97

± 0.18 mm) almost half the length of the inter — orbital space (ISµ = 8.46 ± 0.45 mm;

OD / ISµ = 0.47 ± 0.00) (Fig. 4.4a). Inter — orbital space almost one — fourths carapace width (IS / CW µ = 0.24). Antero — lateral teeth six in number, first slightly notched, sixth greater than two times the length of the preceding five and laterally directed; teeth separated by narrow notches (Fig. 4.4d). Transverse ridges of carapace faintly granular, two proto — gastric, one meso — gastric, a medially divided meta —

76 (III) (Iv)

OD

(c) (v) (d)

(i) Narrow V-shaped notch separating sub-median and lateral frontal teeth (ii) Rounded median frontal teeth (iii) First antero-lateral tooth slightly notched (iv) Narrow notches separate the antero-lateral teeth (v) Last antero-lateral tooth largest and laterally directed

Fig. 4.4. Charybdis (Charybdis) philippinensis Ward, 1941 (a) Dorsal surface of carapace (Photograph), (b) Ventral surface of carapace (Photograph), (c) Frontal margin of carapace (Camera lucida diagram), (d) Antero — lateral margin of carapace (Camera lucida diagram) gastric and epibranchials along the level of the last antero — lateral teeth; no transverse

ridges beyond the level of the last antero — lateral teeth (Fig. 4.4a). Cardiac region

devoid of tuberosity (Fig. 4.4a). Chelipeds subequal (LC pt = 36.30 f 3.96 mm; RC g

= 40.08 ± 2.15 mm), smooth; merus with two well — developed spines and five to six

granules on its anterior margin; manus tumid, five — costate, with five spines on upper

surface, two blunt spines on distal margin immediately posterior to articulation with

dactylus; dactylus smooth, strong, slender with curved tips, longer than manus (Fig.

4.4b). Propodus of natatory legs smooth, merus bears prominent spine on its postero —

distal margin. Penultimate abdominal segment broader than long, its sides parallel

over less than half their lengths and then gradually converge distally (Fig. 4.4b).

Comparison between the new species and its closest congener, C. philippinensis revealed that the new species possessed "bluntly triangular frontal

teeth; wide U — shaped notch separating sub — median and lateral frontal teeth" as

compared to "rounded median frontal teeth; narrow V — shaped notch separating sub —

median and lateral frontal teeth" of the latter species. In addition, antero — lateral teeth

of C. goaensis were separated by "wide notches" as compared to "narrow" ones of C. philippinensis. Moreover, C. goaensis differed from the latter species by the presence

of "granular tuberosity" on cardiac region. A taxonomic key to forty three species of

the subgenus Charybdis is provided below to justify identification of a new portunid

crab species from Goa, west coast of India.

77 4.3.6. Key to the species of swimming crabs of the sub — genus Charybdis

1. Transverse ridges on carapace posterior to the last antero — lateral tooth

absent .. .2.

- Transverse ridges on carapace posterior to the last antero — lateral tooth

present 28.

2. Antero — lateral margins of carapace appear to be divided into five teeth as

first two teeth are grown together except for distal end rendering "bifurcated

first tooth" appearance Charybdis (Charybdis) heterodon Nobili, 1906.

Antero — lateral margins of carapace appear to be divided into seven teeth due

to division of first tooth into larger (anterior) and smaller (posterior)

teeth ..Charybdis (Charybdis) demani Leene, 1937.

Antero — lateral margins of carapace divided into six teeth .3.

3. Two spines on the anterior margin of merus of cheliped. First antero — lateral

tooth square — cut to obtusely triangular; last tooth greater than twice in length

than any of the preceding teeth, and projected laterally. Posterior margin of

propodus of natatory (last) leg smooth .4.

Three spines on the anterior margin of merus of cheliped. Firt antero — lateral

tooth usually smaller than succeeding ones, may be truncated; length of last

antero — lateral tooth may or may not exceed that of preceding spine, may be

projected laterally. Posterior margin of propodus of natatory leg serrated or

smooth 6.

- Three to five spines on the anterior margin of merus of cheliped. First antero —

lateral tooth smallest, may or may not be truncated, last one largest and

78 forwardly, laterally or backwardly directed. Posterior margin of propodus of

natatory leg serrated 22.

4. First antero — lateral tooth obtusely triangular. Two spines on upper surface of

manus of cheliped ..Charybdis (Charybdis) anisodon (De Haan, 1833).

First antero — lateral tooth distinctly bifid. Three spines on upper surface of

manus of cheliped Charybdis (Charybdis) ihlei Leene & Buitendijk, 1949.

First antero — lateral tooth slightly notched. Five spines on upper surface of

manus of cheliped 5.

5. Median frontal teeth bluntly triangular, sub — medians and laterals separated

by wide U — shaped notch. Antero — lateral teeth separated by wide notches.

Prominent granular tuberosity on cardiac region Charybdis (Charybdis)

goaensis sp. nov.

Median frontal teeth rounded, sub — medians and laterals separated by narrow

V — shaped notch. Antero — lateral teeth separated by narrow notches. Lack of

tuberosity on cardiac region.. ..Charybdis (Charybdis) philippinensis Ward,

1941.

6. Upper surface of manus of cheliped with four spines .7

- Upper surface of manus of cheliped with five spines 10.

7. First antero — lateral tooth spiniform, second rudimentary, attached to posterior

margin of first 8.

- First antero — lateral tooth notched or blunt, second not rudimentary 9.

79

8. Inner surface of manus of cheliped smooth; merus of fifth leg proportionally

longer .Charybdis (Charybdis) orientalis Dana, 1852.

Inner surface of manus of cheliped strongly granular; merus of fifth leg

relatively shorter Charybdis (Charybdis) hawaiensis Edmondson, 1954.

9. First antero — lateral tooth distinctly bifid, second and fifth teeth larger than

first, last tooth produced laterally...... Charybdis (Charybdis) feriata

(Linnaeus, 1758).

First two antero — lateral teeth blunt, second and fifth teeth smallest, last tooth

not produced laterally ..Charybdis (Charybdis) padadiana Ward, 1941.

10.Manus of cheliped bears five spines on upper surface, two on distal margin

tuberculated 11.

- All five spines on upper surface of manus of cheliped well

developed .12.

11.Margin of median frontal teeth well in advance of that of sub — medians;

presence of faint transverse ridges on frontal region of

carapace Charybdis (Charybdis) rosaea (Hombron & Jacquinot, 1846).

Margin of median frontal teeth slightly in advance of that of sub — medians; no

transverse ridge on frontal region Charybdis (Charybdis) annulata

(Fabricius, 1798).

12.Posterior margin of carpus of natatory leg bears spine 13.

- Posterior margin of carpus of natatory leg smooth, lacks spine 15.

13.Last antero — lateral spine elongate, projects beyond preceding spine ..14.

80 - Last antero — lateral spine smallest, does not project beyond preceding

spine .Charybdis (Charybdis) vannamei Ward, 1941.

14.Median frontal teeth sharp, triangular, projecting beyond narrow sub —

medians; posterior margin of propodus of natatory leg

denticulated Charybdis (Charybdis) hellerii (A. Milne Edwards, 1867).

Median and sub — median frontal teeth broadly rounded, medians project

slightly in advance of sub — medians; posterior margin of propodus of natatory

leg smooth Charybdis (Charybdis) spinifera (Miers, 1884).

15.Posterior margin of propodus of natatory leg serrated ..16.

- Posterior margin of propodus of natatory leg smooth 21.

16.All frontal teeth sharply triangular Charybdis

(Charybdis) acuta (A. Milne Edwards, 1869).

- All frontal teeth rounded 17.

- Frontal teeth either bluntly triangular or rounded .18.

17.Median frontal teeth slightly in advance of sub — medians, laterals least

advanced Charybdis (Charybdis) incisa Rathbun, 1923a.

- Median frontal teeth well in advance of sub — medians, laterals slightly in

advance of sub — medians Charybdis (Charybdis) javanensis Zarenkov,

1970.

18.Frontal teeth bluntly triangular; median teeth blunter than sub — medians and

laterals 19.

81 - Median and sub — median frontal teeth broadly rounded, laterals broadly

triangular 20.

19. Sub — median frontal teeth separated from the laterals by deep V — shaped

incision. Transverse ridges of carapace faintly granular Charybdis

(Charybdis) lucifera (Fabricius, 1798).

- Sub — median frontal teeth separated from the laterals by wide and deep

fissure. Transverse ridges of carapace distinctly granular Charybdis

(Charybdis) amboinensis Leene, 1938.

20. Cardiac region of carapace with corrugated surface; first antero — lateral tooth

markedly truncated; outer surface of manus strongly squamiform throughout;

G1 of male short, stout and sharply curved, absence of upper inner spines

proximal to lip of G1 Charybdis (Charybdis) yaldwyni Rees &

Stephenson, 1966.

- Cardiac region of carapace typically granulated; first antero — lateral tooth

slightly truncated; outer surface of manus coarsely granular on upper half and

squamiform on lower half; G1 of male long, slender with sinuous margin,

presence of inconspicuous row of well — spaced spinules proximal to

lip Charybdis (Charybdis) jaubertensis Rathbun, 1924.

21. Manus of cheliped with well developed spines; penultimate segment of male

abdomen with convex lateral borders; last antero — lateral tooth smallest and

spiniform, not projecting beyond preceding tooth Charybdis

(Charybdis) japonica (A. Milne Edwards, 1861).

- Manus of cheliped with poorly developed spines; penultimate segment of male

abdomen with lateral margins parallel for proximal half; last antero — lateral

82 tooth elongated, projecting laterally beyond preceding tooth Charybdis

(Charybdis) affinis Dana, 1852.

22.Carapace pilose and narrow, CW / CL ratio being 1.41, ridges of carapace

distinct; lower surface of manus with squamiform marking Charybdis

(Charybdis) miles (De Haan, 1835). - Carapace generally lacking pilosity, relatively broad, CW / CL ratio being 1.49 — 1.67, ridges of carapace faint; squamiform marking on manus, if

present may be faint .23.

23.Ridges on dorsal surface of carapace finely granular, mesogastric ridges distinct; third to fifth antero — lateral teeth with straight edges, others with

sinuous edges...... Charybdis (Charybdis) meteor Spiridonov & Tiirkay, 2001. Ridges on carapace finely or distinctly granular; all antero — lateral teeth with sinuous edges 24.

24.Transverse ridges of carapace distinctly granular; inner sub — orbital lobe broadly triangular; manus massive with faint squamiform marking on lower

surface of outer side 25. - Transverse ridges of carapace finely granular; inner sub — orbital lobe sharply triangular or styliform; manus slender, devoid of squamiform marking on

outer side .26.

25.Carina on inner as well as outer surface of manus in the form of conspicuous ridges; movable finger long, slender and deeply grooved; postern — distal

margin of propodus of natatory leg smooth Charybdis

(Charybdis) rufodactylus Stephenson & Rees, 1968.

83 - Carina on inner as well as outer surface of manus in the form of rounded

ridges; movable finger short, relatively stout and shallow grooved; two

spinules on postero — distal margin of propodus of natatory leg Charybdis

(Charybdis) sagamiensis Parisi, 1916.

26. Inner supra — orbital lobe styliform, outer sharpened; dactyli of second to

fourth pereiopods thin, curved with lancet — shaped tip Charybdis

(Charybdis) acutidens Tiirkay, 1986.

Inner supra — orbital lobe broadly to sharply triangular; dactyli of second to

fourth pereiopods moderately wide, slightly curved .27.

27. Median frontal teeth acutely rounded; lateral frontals with outer edge convex;

basal antennal joint bearing distinct granules; lip of G1 of male tapering with

conspicuous spinules evenly distributed along edges; lateral spines not closely

set, extending to about one — fifths of neck . Charybdis

(Charybdis) riversandersoni Alcock, 1899.

- Median frontal teeth sharply granular; lateral frontals with straight edges;

basal antennal joint bearing fused granules; lip of G1 of male elongated and

bent downward, spinules concentrated on abdominal edge near tip; lateral

spines closely set, extending to one — fourths of neck Charybdis

(Charybdis) crosnieri Spiridonov & Tftrkay, 2001.

28. Transverse ridge on cardiac region interrupted .29.

- Transverse ridge on cardiac region not interrupted 36.

29. Small — sized species; mesobranchial region of carapace with two pairs of

transverse ridges ..30.

84 - Species of a good size; mesobranchial region of carapace with two to three

pairs of transverse ridges . 33

30. Second antero — lateral tooth of carapace smaller than the first, and attached to

its posterior margin Charybdis (Charybdis) rathbuni Leene, 1938.

Second antero — lateral tooth as large as the first, not attached to the posterior

margin of the latter 31.

31. Sixth antero — lateral tooth largest, spiniform and laterally projected ...32.

- Sixth antero — lateral tooth distinctly smaller than preceding tooth, laterally

projected Charybdis (Charybdis) salehensis Leene, 1938.

32. All frontal teeth acuminate at tip. Sub — distal portion of male gonopod

separated as distinct lobe from tip Charybdis (Charybdis) variegata

(Fabricius, 1798).

Only lateral frontal teeth acuminate at tip. Sub — distal portion of male

gonopod not distinct from tip .Charybdis (Charybdis) brevispinosa

Leene, 1937.

33. Dorsal surface of carapace even and uniformly covered with soft tomentum.

Penultimate segment of male abdomen not convex on lateral margins 34.

- Dorsal surface of carapace somewhat uneven, unevenly tomentose.

Penultimate segment of male abdomen convex on lateral

margins . Charybdis (Charybdis) granulata (De Haan, 1833).

34. First antero — lateral tooth of carapace truncated at tip. Frontal ridge of

carapace absent Charybdis (Charybdis) natator (Herbst, 1789).

85 - First antero — lateral tooth not truncated. Frontal ridge present 35.

35. Carapace with high relief and prominent transverse granular ridges. Granules

on chelipeds very prominent and projecting Charybdis

(Charybdis) beauforti Leene & Buitendijk, 1949.

Carapace with low relief and less prominent transverse granular ridges.

Granules on chelipeds less prominent Charybdis (Charybdis)

seychellensis Crosnier, 1984.

36. Mesobranchial region of carapace with two pairs of transverse ridges. Two

spines on anterior margin of merus of cheliped Charybdis

(Charybdis) curtilobus (Stephenson & Rees, 1967b).

- Mesobranchial region of carapace with one pair of transverse ridges. Three

spines on anterior margin of merus of cheliped Charybdis

(Charybdis) cookei Rathbun, 1923b.

- No transverse ridges on mesobranchial region ...37.

37. Carapace about four — fifths as long as broad; two spines on palm of cheliped.

Lateral margins of sixth abdominal segment of male

curved Charybdis (Charybdis) rostrata (A Milne Edwards, 1861).

Carapace about two — thirds as long as broad; three spines on palm of

cheliped. Lateral margins of sixth abdominal segment of male parallel for half

their lengths Charybdis (Charybdis) callianasa (Herbst, 1801).

86 4.4. Conclusion

In view of the above description and comparison of morphological characters with its congeners, C. goaensis is found to be new species to science (Padate et al., 2010b).

The distinguishing characters of the new species are as follows:

1. "Median frontal teeth bluntly triangular",

2. "Sub — median and lateral frontal teeth separated by wide, U — shaped notch",

3. "First antero — lateral tooth slightly notched"

4. "Last tooth of antero — lateral margin greater than twice the length of the

preceding five and laterally directed",

5. "Antero — lateral teeth separated by wide notches",

6. "Prominent granular tuberosity on cardiac region",

7. "Two spines on the anterior margin of merus of cheliped",

8. "Five spines on upper surface of manus of cheliped, two on distal margin

remain tuberculated",

9. "Membrane on tip of G1 of male commences shortly behind the tip; row of

sparsely placed bristles on outer margin starts at some distance from the tip,

but anterior to the above membrane; row of four to five bristles very sparsely

placed on inner margin commencing distantly from distal tip and terminating

prior to outer row of bristles; bristles on abdominal margin randomly and very

widely placed."

87 Chapter 5. A new record of Caesio cuning outside its known geographical array 5.1. Introduction

The zoo — geographical range of a species refers to the distribution of a species, either horizontal or vertical, largely determined by the pervasiveness of certain environmental anomalies and biotic components those favour establishment of the species, and whose boundaries are defined by gradual dilution or abrupt termination of these components (Padate et al., 2010a). In the marine environment, physical factors such as temperature, salinity, dissolved oxygen and nutrient levels may tend to regulate the occurrence and distribution of a species through availability of adequate food and substratum, as well as regulation of biological processes such as recruitment, establishment and reproduction. Coastal ecosystems being dynamic are rich in diverse assemblages with estimates of about 80 % of marine fauna inhabiting these areas (Anon., 1989; Ray, 1988, 1991) with highly overlapping boundaries those are influenced by complex relationships among varying niches in the marine environment. These coastal areas also support rare fish species those occur seasonally or occasionally in very low numbers (Engel and Kvitek, 1998). Among these, species belonging to two orders, namely Perciformes (perch — like fishes) and Scorpaeniformes (scorpion fishes, rock fishes and sting fishes) dominate the fish assemblages (Froese and Pauly, 2008). Further among the perch — like fishes, the fusiliers belonging to the family Caesionidae are particularly unique with regards to evolutionary trends those display transition from benthic carnivorous to semi — pelagic planktivorous habit (Carpenter, 1988).

5.2. Literature review

Published literature pertaining to the taxonomy of the Family Caesionidae reveals limited works by various taxonomists across the Indo — West Pacific region

88 (Masuda et al., 1975; Schroeder, 1980; Shen, 1984; Gloerfelt — Tarp and Kailola,

1984). However, Carpenter (1987, 1988) provided exhaustive descriptions of twenty extant fusilier species along with phylogenetic classification. Further, among the species described in the above literature, Caesio cuning (Bloch, 1791) is known to be distributed from southern Japan to northern Australia and from Vanuatu to the Bay of

Bengal (Carpenter, 1988).

In view of the above, this chapter describes the new record of occurrence of the rare fusilier species, C. cuning found outside its geographical locale in the coastal waters of Goa along the west coast of India (Padate et al., 2010a).

5.3. Taxonomy

5.3.1. Family Caesionidae

The diagnostic characters of Caesionidae are "body slender, fusiform, elongate

(resembling pelagic fishes) and brightly coloured with stripes and patches; unique jaw morphology, wherein the ascending maxillary process is completely separated from the pre — maxilla; highly protrusible upper jaw; reduced dentition."

The above morphological characters are an adaptation towards planktivorous feeding habit. Further, they exhibit synchronous schooling habit that facilitates their semi — pelagic mode of locomotion in the coastal waters. Contrastingly, in other related perciform fishes such as the benthic carnivores, particularly snappers (family

Lutjanidae), the ascending maxillary process is confluent with the pre — maxilla

(Carpenter, 1988), and the dentition consists of strong canines to facilitate grasping of prey.

89 5.3.2. Genus Caesio Lacepecle, 1801 (Carpenter, 1988)

Caesio Lacepede, 1801, pp. 85 (type species: Caesio caerulaurea Lacepede, 1801, by subsequent designation (Bleeker, 1876))

Fin formula: D X, 13 — 16; A III, 10 —13; P 17 — 23; V I, 5; C (procurrent) 9 —10

"Body high to fusiform, elongate and moderately compressed. A single post — maxillary process; posterior end of maxilla blunt, its greatest depth posterior to end of pre — maxilla; small conical teeth on jaws, vomer and palatines; interorbital space convex; margin of opercle with a pronounced dorso — posterior flap."

"Scales weakly ctenoid; scales present on dorsal and anal fins, those in lateral line 45 — 67; 9 — 13 scale rows on the upper portion of caudal peduncle, 12 — 17 on the lower one; 7 — 11 scale rows between lateral line and dorsal fin origin, 14 — 20

scale rows between lateral line and anal fin origin; supra — temporal band of scales

distinct, confluent at dorsal midline or interrupted by a thin scaleless zone."

"Predorsal configuration 0/0/0 + 2/1 + 1/. 10 — 14 epipleural ribs, without

flattened projections on first or second epipleural. Anterior profile of first anal pterygiophore strongly convex distally."

"Colour: lateral sides may possess longitudinal stripes; caudal fin either

devoid of markings, with a blackish blotch on tips of lobes, or with a longitudinal

blackish streak in the middle of each lobe."

5.3.3. Caesio tuning (Bloch, 1791)

5.3.3a. Material examined

Two unsexed specimens; RP — 1, 6.08 cm SL and RP — 2, 5.21 cm SL (Co —

ordinates: 15°30'43.1"N, 73°45'40.1"E to 15 °32'28.1"N, 73°45'9.4"E; water depth:

11 — 14 m). Two unsexed specimens; RP — 3, 6.89 cm SL and RP — 4, 6.11 cm SL (Co

90 — ordinates: 15°29'43.2"N, 73°44'39.7"E to 15°33'15.2"N, 73°42'41.6"E; water depth: 6 — 7 m).

Details of morphometric measurements of fish specimens along with their mean and standard deviation are provided in Table 5.1. Reference voucher sample is deposited at the Marine Biology laboratory, Department of Marine Sciences, Goa

University.

5.3.3b. Description

Fin formula: D X, 15; A III, 11; P 17; V I, 5; C (procurrent) 9 —10

Dorsal region of body from tip of snout to the anterior spinous portion of dorsal fin greyish yellow, soft dorsal fin bright yellow, mid — lateral portion of body pale white, a black blotch on pectoral fin axil (Fig. 5.1 a); caudal fin yellow, lacking prominent blackish markings (Fig. 5.1b); ventral region of body including the cheeks, pectoral, pelvic, anal fins and the lower portion of caudal peduncle, light pink (Fig.

5.1c). Body fairly deep (body depth approximately three times in standard length,

almost equal to head length) and compressed. A single post — maxillary process; small

conical teeth in jaws, vomer and palatines. Supra — temporal rows of scales on either

side of head confluent at midline (Fig. 5.2); four to five scale rows on cheek; 48 — 51

scales in lateral line; eight scale rows between lateral line and the origin of dorsal fin;

fifteen scale rows between lateral line and the origin of anal fin. Dorsal fin

continuous, with ten spines and fifteen soft rays, scales cover approximately half the

greatest height of dorsal fin (Fig. 5.3); anal fin with three spines and eleven soft rays

(Fig. 5.4); spines on both the dorsal and anal fins interconnected by thin membrane;

pectoral fins with seventeen soft rays; pelvic fins with one spine and five soft rays;

caudal fin strongly forked, with nine to ten procurrent rays.

91 Table 5.1. Details of morphometric measurements (in cm) of Caesio cuning specimens (N = 04) collected off Goa during the present study

Sr. Morphological Specimen ft ± S.D. No. parameter RP - 1 RP - 2 RP - 3 RP - 4

1. Total length 8.48 7.49 10.05 8.56 8.65 ± 1.06 2. Fork length 6.99 6.57 8.38 7.42 7.34 ± 0.78 3. Standard length 6.08 5.21 6.89 6.11 6.07 ± 0.69 4. Body depth 2.34 1.91 2.64 2.27 2.29 ± 0.30 5. Pre - dorsal Length 2.28 2.07 2.53 2.32 2.30 ± 0.19 6. Pre - anal Length 4.18 NA 4.84 4.33 4.45 1 0.35 7. Head length 2.02 NA 2.36 2.13 2.17 ± 0.17 8. Snout length 0.39 0.42 0.44 0.38 0.41 ± 0.03 9. Eye diameter 0.74 0.61 0.88 0.87 0.78 ± 0.13

RP - site of sample collection in the vicinity of the marine vessel River Princess grounded off Goa coast Fig. 5.1 a. Distinguishing characters and colour of fresh specimen of Caesio cuning (Photograph)

Fig. 5.1b. Caudal fin of Caesio cuning indicating characteristic yellow colouration (Photograph)

Fig. 5.1c. Ventral portion of Caesio cuning indicating characteristic pink colouration (Photograph) Fig. 5.2. Supra — temporal band of scales in Caesio tuning (a) Photograph, (b) Diagrammatic representation (Carpenter, 1988) Fig. 5.3. Dorsal fin of Caesio cuning indicating scaled portion and characteristic fin count (Photograph)

Fig. 5.4. Anal fin of Caesio cuning indicating characteristic fin count (photograph) Morphometric data of the present specimens (Table 5.1) was compared with the existing literature. Fishbase (Froese and Pauly, 2008) revealed that C. culling grows up to a size of 60 cm Total length (TL). However, the present specimens (N =

04) ranged, between 7.49 and 10.05 cm TL = 8.65 ± 1.06 cm), thus confirming the capture of juvenile / young individuals of this species.

5.4. Discussion

The existing data on the geographical distribution of Caesio cuning (Fig. 5.5) suggests that this species inhabits the Indo — Pacific region from Gulf of Mannar to

Vanuatu Islands in the South Pacific (Carpenter, 1988; Froese and Pauly, 2008). The

above regions fall within the tropical zone and are characterized by warm, saline, and nutrient — rich waters. Further, these regions harbour diverse habitats such as coral reefs, mudflats, sea — grass beds, thereby augment to the high productivity of these

waters. Carpenter (1988) attributed the geographical distribution of C. cuning to the

availability of appropriate habitat and adequate ecological parameters those favour the

occurrence of this species.

Published literature (Talwar and Kacker, 1984) pertaining to the rare fishes of

Indian coast provide a short description of C. cuning along with short note on its

occurrence and distribution. In Indian territory, this species is known to occur only

along the Andaman coast which incidentally falls within known zoo — geographical

range of this species (Talwar and Kacker, 1984). Although rare in occurrence and

distribution, this species finds mention in the above work due to its commercial

importance.

However, Fishbase (Froese and Pauly, 2008) indicates the possible expansion

of occurrence of this species to the adjacent waters due to changes in climate or land

92 ❑ — Known zoo — geographical range of Caesio cuning a— Possibility of occurrence of Caesio cuning GOA — Site of present collection Fig. 5.5. Map indicating the worldwide distribution of Caesio cuning use patterns those were responsible for the development of ambient conditions, thereby favouring the occurrence of this species. Published literature along Goa coast

(Prabhu and Dhawan, 1974; Ansari et al., 1995, 2003) provides information on the demersal fish community structure as well as the fishery potential of this region.

However, efforts to obtain information pertaining to the entire demersal faunal community have resulted in scanty information as evident from the reporting of the most dominant or common species. C. cuning being a rare species does not find mention in available literature from the present study area.

In view of the above, our observations primarily focus on the occurrence of rare species along Goa coast. Subsequently, the present investigation led to the reporting of C. cuning for the first time off Goa and along the entire west coast of

India. It has been well documented earlier (Carpenter, 1988) that this species and its congeners occur in coastal reef areas. The present specimens were obtained from bottom trawl catch in the vicinity of the grounded ore — carrier, MV River Princess off the Candolim — Sinquerim shore. The grounded MV River Princess off Goa coast in

2000 (Ingole et al., 2006) might have created an artificial reef — like habitat, thereby attracting several reef inhabitants from adjacent reefs (Qasim and Wafar, 1979;

Rodrigues et al., 1998) and further facilitated recruitment of their larvae and juvenile forms (Arena et al., 2007). The observations made on size of collected individuals (..t

= 8.65 t 1.06 cm) suggested capture of young stages. Published literature (Qasim and

Wafar, 1979; Rodrigues et al., 1998) indicates the presence of partially submerged rocky reefs off the Aguada Hill, suggesting suitability of habitat. Moreover, the above region contains nutrient — rich turbid waters due to transport of suspended material

from the adjacent Mandovi estuary and Aguada bay (Ansari et al., 1995). Kessarkar et al. (2009) reported seasonal peaks in suspended particulate matter (SPM)

93 concentration, 20 and 19 mg.r 1 during June — September and February — April, respectively from the Mandovi estuary and attributed these to interactions between strong seasonal winds, wind — induced waves and tidal currents in the estuary. It is probable that wind — induced waves, tidal currents along with the resultant turbidity in the water column might have induced the migration of C. cuning juveniles towards the bay — estuarine waters. Carpenter (1988) suggested that C. cuning prefers turbid waters, thus corroborating with the present observations. Further, Leis and Carson —

Ewart (2003) suggest that pelagic larvae of certain reef fish orient their movement in response to solar or reef — based cues. Caesio cuning inhabits the eastern Indian

Ocean and Western Pacific Ocean regions (Carpenter, 1988) those influenced by hydrographic parameters (temperature, salinity, dissolved oxygen) characteristic of tropical oceans (Moore, 1972). Hydrographic parameters such as temperature (27.5 —

28.7 °C), salinity (35.1 psu) and DO (180 — 190 gM) were observed to be in similar range and indicated clear seasonal variations at 10 m depth (Fig. 5.6), however the values were found to be comparatively higher at the time of collection of C. cuning.

Similar studies on C. cuning ecology from the Seribu Island, Indonesia (Tati et al.,

1998) reported marginally different values for temperature (28.7 — 29.5 °C), salinity

(32.55 — 32.73 psu) and DO (3.85 — 4.25 mg.1' 1). Further, the study area experiences seasonal upwelling (Ansari et al., 1995) that brings the cold, nutrient — rich, sub — surface water to the surface. The above phenomenon sustains luxurious phytoplankton growth and renders the region highly fertile in terms of primary productivity (13.12 —

14.21 mg C.m-3.111 ; Krishna Kumari et al., 2002). Such high productivity sustains high zooplankton biomass (Qasim and Sen Gupta, 1981). C. cuning preferably feeds on mid — water zooplankton (Carpenter, 1988), and it is possible that schools of this

94 36 -

3 S. 34

32 xi

30

200

& 150

100 0 50 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Month

Fig. 5.6. Seasonal variations in hydrographic parameters measured at surface and 10 m depth during 2007 species might have migrated to the above region to forage upon the abundant zooplankton.

Further, the intensive sampling effort covering wider coastal region through use of modem fishing vessels and gear those provided access to submerged rocky patches in coastal inshore embayment may have enabled entrapping this species.

However, it must be noted that in the present study, only four specimens were obtained in the light of the trawl hauls (N = 95) with a core fishing effort of one hundred and fifty six hours over a period of three years excluding monsoonal periods

(June — August). Moreover, only two individuals each were obtained from two hauls with a highly diverse faunal composition. Carpenter (1988) suggested a shoaling behaviour among the caesionids, similar to those in other planktivorous fishes.

However, no large shoals were observed during the present study. This probably

could be due to segregation of these individuals from the larger shoal to be trapped in

the trawl net or might have not been able to establish as a large shoal. Fishbase

(Froese and Pauly, 2008) revealed that this species inhabits rocky substrates in coastal

waters up to 60 m depth and forms large shoals in mid — water. It is apparent from the

above that the low depth 10 m) along with the habitat complexity i.e. silt — clayey

substratum interspersed with submerged rock outcrops and patch reefs might have

probably obstructed the formation of a large shoal. Moreover, the study area is a

potential fishing ground and has been intensively exploited by demersal trawlers

perennially with the exception of the seventy five — day monsoon ban ever since the

initiation of mechanized fishing and its further expansion during the subsequent

decades. Such long — term intensive trawling activity may lead to changes in the

structure and composition of the resident demersal communities and benefit other

95 species groups as reported earlier from the Gulf of Thailand (Longhurst and Pauly,

1987).

5.5. Conclusion

Our observations suggest the first new record of Caesio curling off Goa along the entire west coast of India outside its known geographical array (Padate et al.,

2010a). The occurrence of this species in low numbers could primarily be due to sampling gear and local bathymetry in the vicinity of the grounded ship that probably reduced the chances of entrapment of this species in the trawl net. Furthermore, it appears that the selection of gear with increased efficiency in such reef dominated habitats might yield more individuals from such areas. It is suggested that further detailed ecological studies need to be taken up to provide better insight into its occurrence and distribution. Moreover, molecular studies of this species using genomic DNA would further enhance the understanding of its population stock

structure and connectivity with existing populations (Carmen and Ablan, 2006).

96 Chapter 6. A new record of Scylla olivacea from Goa - A comparative diagnosis 6.1. Introduction

Mud crabs of the genus Scylla (de Haan, 1833) are one of the most common inhabitants of mangrove — vegetated estuarine regions throughout the Indo — Pacific region (Stephenson, 1962). These crabs are highly valued in the international market because of their large size and meat quality (Marichamy and Rajapackiam, 2001).

Consequently, efforts have been made to elevate crab production through mariculture of these crabs (Marichamy and Rajapackiam, 2001). However, it is suggested that the uncertainty in genetic relationship and dearth of taxonomic details among the mud crabs render the implementation of fisheries management regulations and development of sustainable aquaculture difficult (BOBP, 1992). Therefore, it is imperative to obtain an insight into the history of taxonomy of the mud crabs belonging to the genus Scylla for improved understanding of the problems associated with identification of these crabs.

6.2. Literature review

Published literature (BOBP, 1992) suggests the occurrence of various species or "morphs" of the crabs belonging to genus Scylla, their taxonomic status under considerable ambiguity (Joel and Raj, 1980) in view of the localized variations in morphology of these crabs (Marichamy and Rajapackiam, 2001). Earlier authors

(Stephenson and Campbell, 1960; Overton et al., 1997) have attributed these variations to the ambient environmental conditions. Taxonomy of mud crabs was

initiated by Forskal (1775). Subsequently, various naturalists identified mud crabs

using orthodox identification methods and reported as many as six "species" or

"varieties" from throughout the Indo — Pacific region (Herbst, 1796; Fabricius, 1798;

Riippell, 1830; De Haan, 1833; Macleay, 1838; H. Milne Edwards, 1834; Dana, 1852;

97 A. Milne Edwards, 1861; Henderson, 1893; Alcock, 1899a; Lanchester, 1900; de Man, 1909; Stebbing, 1910; Kemp, 1915; Gravely, 1927; Pearse, 1932; Shen, 1932; Chopra and Das, 1937; Pannikar and Aiyar, 1937; Estampador, 1949; Barnard, 1950;

Pillai, 1951; Serene, 1952; Naidu, 1953; Chacko et al., 1953; Chacko, 1956; Chhapgar, 1957; Stephenson and Campbell, 1960; Crosnier, 1962; Ong, 1964; Holthuis, 1978; Joel and Raj, 1980; Kakati, 1980; Radhakrishnan and Samuel, 1982; Fushimi, 1983; Prasad, 1987; Chen, 1989; Oshiro, 1991; Kathirvel and Srinivasagam, 1992; Fuseya and Watanabe, 1995; Watanabe and Fuseya, 1997). In recent years, molecular techniques based on phylogenetic distances are being employed to ascertain species identity and secondly to determine phylogenetic linkages between different populations of a species. Fuseya and Watanabe (1996) carried out genetic variability studies at three loci in the mud crabs and differentiated three species namely Scylla serrata, S. tranquebarica and S. oceanica. However, S.

oceanica was considered to be synonymous with S. serrata and S. paramamosain

(Keenan et al., 1998) and S. tranquebarica (Marichamy and Rajapackiam, 2001).

Overton et al. (1997) employed Canonical Variate Analysis, a statistical approach based on morphometric measurements and meristic counts to facilitate easy differentiation between varieties within a single species those are geographically

separated. Consequently, they reported two forms of S. serrata namely "black" and "white". However, the lack of availability of type specimens has resulted in considerable confusion regarding the taxonomic nomenclature of these crabs coupled with vaguely defined morphological characters those render differentiation among

various species or varieties of Scylla extremely difficult (Keenan et al., 1998). Subsequently, in order to resolve the issue of mud crab species identification, Keenan et al. (1998) used two novel genetic methods, allozyme electrophoresis and

98 sequencing of two mitochondrial DNA genes (cytochrome oxidase 1 and 16S RNA).

These methods revealed four distinct species (S. serrata (Forskal, 1775), S. olivacea

(Herbst, 1796), S. tranquebarica (Fabricius, 1798) and S. paramamosain Estampador,

1949)), and based on the above analysis, a systematic approach comprising measurement of twenty four morphological parameters of mud crabs and subsequent derivation of twenty seven morphometric ratios was adopted to characterize morphometric data to enable field identification and accordingly a key to the existing species of Scylla was formulated as follows.

1. Carpus of chelipeds with two obvious spines on distal half of outer margin...2.

— Carpus of chelipeds without two obvious spines on distal half of outer

margin ...3.

2. Frontal lobe spines high (mean height approximately 0.06 times frontal width

measured between medial orbital sutures), bluntly pointed with tendency to

concave margins and rounded interspaces. Antero — lateral carapace spines

narrow, with outer margin straight or slightly concave. Chelipeds and legs all

with polygonal patterning for both sexes and on abdomen of female

only .Scylla serrata.

— Frontal lobe spines of moderate height (mean height approximately 0.04 times

frontal width measured between medial orbital sutures), blunted with rounded

interspaces. Antero — lateral carapace spines broad, with outer margin convex.

Polygonal patterning weak on chelipeds and first two pairs of legs; last two

pairs of legs with stronger patterning for both sexes; patterning variable on

abdomen of female, absent on male Scylla tranquebarica.

99 3. Frontal lobe spines high (mean height approximately 0.06 times frontal width measured between medial orbital sutures), typically triangular with straight margins and angular interspaces. Palm of cheliped with a pair of distinct spines on dorsal margin behind insertion of the dactyl, followed by ridges running posteriorly. Chelipeds and legs with weak polygonal patterning for

both sexes Scylla paramamosain. — Frontal lobe spines low (mean height approximately 0.03 times frontal width measured between medial orbital sutures), rounded with shallow interspaces. Palm of cheliped usually with a pair of blunt prominences on dorsal margin behind insertion of the dactyl, inner larger than outer; may be spinous in juveniles and young adults. Chelipeds, legs and abdomen all without obvious

polygonal patterning for both sexes Scylla olivacea.

Sugama and Hutapea (1999) successfully attempted delineation of three species of Scylla collected from various localities in Indonesia by allozyme electrophoresis and Principal Component Analysis (PCA). Imai et al. (2004) used genetic markers such as ITS — 1 gene of nuclear rDNA and mitochondrial 16S rDNA to identify four Scylla species. Research on the taxonomy of mud crabs in India was initiated in 1798 when

Fabricius described S. tranquebarica from the southeast coast of India. Subsequent efforts to establish the identity of various "species", "morphs" or "varieties" using orthodox identification techniques (Alcock, 1899a; de Man, 1909; Kemp, 1915; Gravely, 1927; Pearse, 1932; Chopra, 1935; Chopra and Das, 1937; Pannikar and

Aiyar, 1937; Pillai, 1951; Naidu, 1953; Chacko et al., 1953; Chhapgar, 1957; Rekha, 1968; Radhakrishnan and Samuel, 1982; Kathirvel and Srinivasagam, 1992)

100 suggested the occurrence of two species namely S. serrata and S. tranquebarica and one variety, S. serrata serrata. Joel and Raj (1980) reported two species namely S. serrata and S. tranquebarica from the Pulicat lake, east coast of India. However,

Keenan et al. (1998) based on their newly defined criteria for species identification, re

— identified these as S. olivacea and S. serrata, respectively. Kakati (1980) and Prasad

(1987) reported only one species namely S. serrata from Karwar, central west coast of

India. Recently, Gopikrishna and Shashi Shekhar (2003) attempted delineation between two mud crabs species (S. serrata and S. tranquebarica) using karyology and

Restriction Fragment Length Polymorphism (RFLP) of 12S and 16S rRNA mitochondrial PCR products and suggested the use of Hind III restriction profile of

16S rRNA mitochondrial PCR product as potential molecular tool for delineating the above species.

The coastal region of Goa, central west coast of India harbours thickly vegetated mangrove habitats among narrow inter — tidal mudflats along the banks of estuaries (Qasim and Sen Gupta, 1981). Mangrove habitats function as nursery grounds for juveniles of mud crabs, and provide food and shelter for adult stages of mud crabs (Walton et al., 2006; Nagelkerken et al., 2008). However, no attempts have been made to address the taxonomy of mud crabs from this region. In view of this, the present study was taken up to assess the diversity in mud crabs those inhabit the coastal waters of Goa. During the observations on the demersal fauna, certain ambiguity in the mud crab identification prompted the initiation of a comparative taxonomic assessment among various existing congeners of the mud crab Scylla.

101 6.3. Taxonomy

Mud crabs obtained from inshore bottom trawls as well as commercial outlets were subjected to phenotypic analyses including morphology, twenty four morphometric measurements following Keenan et al. (1998) and twenty seven morphometric ratios were derived from the above parameters. However, morphological characters of the male gonopod (G1) such as the form of apex and margins on either sides of apex were not considered for the present exercise owing to very subtle differences in this structure among the congeners.

Terminology used in the morphological description of mud crabs follows

Keenan et al. (1998). The following abbreviations are used: CW — Carapace width;

ICW — Internal carapace width; LSH = (CW — ICW) / 2; 8CW — Carapace width at spine 8; PWC — Posterior width of carapace; CL — Carapace length; FMSH — Frontal median spine height; FW — Carapace frontal width; DFMS — Distance between frontal median spines; DFLS — Distance between frontal lateral spines; SW — Sternal width;

AW — Abdominal width; PL — Propodus length; DL — Dactyl length; PW — Propodus width; PD — Propodus depth; IPS — Inner propodus spine; OPS — Outer propodus spine; ICS — Inner carpus spine; OCS — Outer carpus spine; ML — Merus length; 5PW

— 5th pereiopod dactyl width; 5PL — 5 th pereiopod dactyl length; 3PML — 3 rd pereiopod merus length.

Phenotypic analyses of the mud crabs involving examination of the colouration on the carapace, chelipeds and legs, measurement of morphometric parameters (N = 24) and calculation of ratios (N = 27) derived from the above parameters are based on the methods suggested by Keenan et al. (1998). The above analyses are listed in Tables 6.1 and 6.2.

102 Table 6.1. List of phenotypic characters and morphometric parameters (N = 24) used in morphometric analyses of mud crabs during the present study

Sr. No. Type of analysis I. Phenotypic analysis 1. Colouration on carapace 2. Presence of polygonal patterning on carapace, abdomen, chelipeds and legs H. Morphological parameters A. Carapace 1. Carapace width (CW) 2. Internal carapace width (ICW) 3. Carapace width at spine 8 (8CW) 4. Carapace length (CL) 5. Posterior width of carapace (PWC) 6. 9th lateral spine height (LSH) 7. Frontal width (FW) 8. Frontal median spine height (FMSH) 9. Distance between frontal median spines (DFMS) 10. Distance between frontal lateral spines (DFLS) 11. Sternum width (SW) 12. Abdomen width (AW) B. Cheliped 1. Dactylus length (DL) 2. Propodus length (PL) 3. Propodus width (PW) 4. Propodus depth (PD) 5. Inner propodus spine (IPS) 6. Outer propodus spine (OPS) 7. Inner carpus spine (ICS) 8. Outer carpus spine (OCS) 9. Merus length (ML) C. Pereiopods (Walking legs) 1. 5th pereiopod dactyl length (5PL) 2. 5th pereiopod dactyl width (5PW) 3. 3rd pereiopod menus length (3PML) Table 6.2. List of morphometric ratios (N = 27) derived for morphometric analysis of mud crabs during the present study

Sr. No. Type of analysis A. Carapace 1. 9th lateral spine height (LSH) / Internal carapace width (ICW), where LSH = (CW — 1CW) / 2 2. Carapace width (CW) / Carapace width at spine 8 (8CW) 3. Carapace length (CL) / Internal carapace width (ICW) 4. Posterior width of carapace (PWC) / Internal carapace width (ICW) 5. Frontal width (FW) / Internal carapace width (ICW) 6. Posterior width of carapace (PWC) / Frontal width (FW) 7. Frontal median spine height (FMSH) / Frontal width (FW) 8. Frontal median spine height (FMSH) / Distance between frontal median spines (DFMS) 9. Distance between frontal median spines (DFMS) / Frontal width (FW) 10. Distance between frontal lateral spines (DFLS) / Frontal width (FW) 11. Distance between frontal median spines (DFMS) / Distance between frontal lateral spines (DFLS) 12. Sternum width (SW) / Internal carapace width (ICW) 13. Abdomen width (AW) / Sternum width (SW) B. Cheliped 1. Propodus length (PL) / Internal carapace width (ICW) 2. Dactylus length (DL) / Propodus length (PL) 3. Propodus width (PW) / Propodus length (PL) 4. Propodus depth (PD) / Propodus length (PL) 5. [Propodus width (PW) x Propodus depth (PD) x 0.7854] / Propodus length (PL) 6. Inner propodus spine (IPS) / Propodus length (PL) 7. Outer propodus spine (OPS) / Propodus length (PL) 8. Inner propodus spine (IPS) / Outer propodus spine (OPS) 9. Inner carpus spine (ICS) / Propodus length (PL) 10. Outer carpus spine (OCS) / Propodus length (PL) 11. Inner carpus spine (ICS) / Outer carpus spine (OCS) 12. Merus length (ML) / Propodus length (PL) C. Pereiopods (Walking legs) 1. 5`" pereiopod dactyl width (5PW) / 5 th pereiopod dactyl length (5PL) 2. 3' pereiopod merus length (3PML) / Internal carapace width (ICW) The above ratios were subjected to Student's "t — test" at three different levels of significance (P < 0.05; < 0.01; < 0.001) to assess whether there existed any significant differences among the mud crabs with respect to their morphology.

Preliminary phenotypic analyses of the mud crabs (N --, 31) based on the above morphological characters revealed two "morphs" (forms) of mud crabs. Subsequently, an attempt was made to identify these following the taxonomic key provided by

Keenan et al. (1998).

6.3.1. Morph 1

6.3.1a. Material examined

One female (CW 14.84 cm), 13 December 2005, Betim, Goa; four males (CW 11.38 cm, CW 9.38 cm, CW 9.01 cm, CW 10.23 cm), 27 September 2007, Panaji market,

Goa; two females (CW 14.84 cm, CW 14.84 cm), 27 September 2007, Mandovi estuary, Goa; one male (CW 9.15 cm), one female (CW 10.28 cm), 16 January 2008,

Panaji market, Goa; one male (CW 9.50 cm), 12 March 2008, Panaji market, Goa; one male (CW 11.20 cm), 15 March 2008, Panaji market, Goa.

6.3.1b. Description

Carapace broader than long, with prominent H — shaped groove present on cardiac region (Fig. 6.1 a,b). Frontal margin of carapace (excluding inner supra — orbital angles) with four bluntly pointed spines with slightly concave margins and separated by inverted — V shaped interspaces, their height approximately 0.06 times frontal width measured between the orbital sutures (Fig. 6.2). Antero — lateral margins of carapace longer than the postero — lateral margins, and divided into nine sharp spines; all spines almost equal in size. Chelipeds massive, smooth (Figs. 6.1a,b),

103 (i) and (ii) Prominent polygonal markings on chelipeds

Fig. 6.1. Scylla serrata (a) Dorsal view of carapace (Photograph), (b) Frontal view of carapace (Photograph)

(i) Inverted V-shaped interspaces (ii) Bluntly triangular frontal spines

Fig. 6.2. Scylla serrata — Frontal margin of carapace (Camera lucida diagram) longer than legs; menus with three spines on its anterior margin and two on the posterior margin; carpus with acute spine at the inner angle and two prominent spines at the outer angle (Fig. 6.3); propodus with a strong spine at the articulation with carpus and two well developed spines on the dorsal margin behind insertion of the dactylus (Fig. 6.3); both dactyli slender with blunt pointed tips. Four pairs of pereiopods, first three pairs ambulatory (slightly thickened for walking) and similar, the fourth pair natatorial (flattened for swimming). Male abdomen bluntly triangular in shape; female abdomen watch — glass shaped (Fig. 6.4). G1 with long distal portion, ends in short, sharply pointed tip, its outer margins of apex slightly convex (Fig. 6.5); chromatophores absent.

Colouration of carapace varied, olive green to dark brown with randomly scattered polygonal patterning (Fig. 6.1 a,b), ventral surface of abdomen cream colour (only female abdomen with polygonal patterning; Fig. 6.4), chelipeds and legs olive green with conspicuous polygonal marking, dactyli of chelipeds and legs light brown (Fig. 6.1a,b).

Based on the above description and subsequent comparison with Keenan et al.

(1998), the above morph unambiguously resembled S. serrata. The details of morphometric measurements (range, means and standard deviation) are provided in Table 6.3.

It is pertinent to note that these crabs share the character "Palm of chelipeds with pair of distinct spines on dorsal margin behind insertion of the dactyl" (Fig. 6.3) with three species (S. serrata, S. tranquebarica and S. paramamosain) described by

Keenan et al. (1998).

104 (i )

(ii)

(i) Well developed propodal spines behind dactyl insertion (ii) Prominent spines on outer margin of carpus

Fig. 6.3. Scylla serrata Fig.6.4. Scylla serrata — Colour — Right cheliped (Photograph) pattern on female abdomen (Photograph)

(b) (c)

(i) Long, broad distal portion of GI (ii) Short and sharply pointed apex (iii) Slightly convex outer margin

Fig. 6.5. Scylla serrata (a) Entire GI , (b) Tip of GI , (c) Tip of G1 (enlarged view) (Camera lucida diagrams) Table 6.3. Range, mean and standard deviation of morphometric ratios of S. serrata (N = 11) and S. olivacea (N = 20) obtained during the present study

Sr. Morpho - S. serrata S. olivacea No. metric ratio (N = 11) (N = 20) Range It ± S.D. Range it ± S.D. 1. FMSH / FW 0.049 - 0.077 0.062 ± 0.009 0.026 - 0.066 0.045 ± 0.012 2. FW / ICW 0.383 - 0.463 0.424 ± 0.024 0.397 - 0.472 0.431 ± 0.019 3. ICS / OCS 0.000 - 5.636 2.155 ± 1.844 0.000 - 0.500 0.101 ± 0.177 4. LSH / ICW 0.015 - 0.044 0.027 ± 0.009 0.016 - 0.047 0.026 ± 0.008 5. CW / 8CW 0.969 - 1.028 1.000 ± 0.017 0.977 - 1.037 1.004 ± 0.016 6. CL / ICW 0.670 - 0.709 0.695 ± 0.012 0.669 - 0.701 0.688 ± 0.009 7. PWC / ICW 0.297 - 0.416 0.353 ± 0.036 0.318 - 0.392 0.356 ± 0.023 8. PWC/FW 0.662 - 1.076 0.838 ± 0.128 0.709 - 0.940 0.829 ± 0.081 9. FMSH / DFMS 0.306 - 0.738 0.515 ± 0.118 0.217 - 0.615 0.371 ± 0.108 10. DFMS/FW 0.100 - 0.161 0.124 ± 0.018 0.094 - 0.141 0.122 ± 0.013 11. DFLS / FW 0.000 - 0.152 0.115 ± 0.040 0.112 - 0.252 0.139 ± 0.029 12. DFMS / DFLS 0.820 - 1.371 0.995 ± 0.143 0.467 - 1.233 0.900 ± 0.152 13. SW / ICW 0.519 - 0.578 0.541 ± 0.016 0.520 - 0.552 0.538 ± 0.010 14. AW / SW 0.580 - 0.930 0.709 ± 0.137 0.580 - 0.902 0.688 ± 0.125 15. PL/ICW 0.610 - 0.888 0.745 ± 0.087 0.000 - 0.878 0.657 ± 0.236 16. DL / PL 0.416 - 0.525 0.466 ± 0.028 0.000 - 0.510 0.421 ± 0.145 17. PW / PL 0.248 - 0.286 0.270 ± 0.014 0.000 - 0.430 0.254 ± 0.095 18. PD/PL 0.372 - 0.448 0.417 ± 0.023 0.000 - 0.485 0.368 ± 0.133 19. (PW x PD x 0.558 - 0.863 0.671 ± 0.090 0.000 - 2.794 0.683 ± 0.555 0.7854) / PL 20. IPS / PL 0.028 - 0.104 0.057 ± 0.022 0.000 - 0.089 0.042 ± 0.025 21. OPS / PL 0.016 - 0.040 0.029 ± 0.008 0.000 - 0.038 0.018 ± 0.011 22. IPS / OPS 1.400 - 3.450 2.128 ± 0.597 0.000 - 5.000 2.553 ± 1.140 23. ICS / PL 0.000 - 0.103 0.025 ± 0.038 0.000 - 0.016 0.004 ± 0.006 24. OCS / PL 0.000 - 0.063 0.019 ± 0.020 0.000 - 0.056 0.023 ± 0.015 25. ML / PL 0.521 - 0.671 0.579 ± 0.037 0.000 - 0.628 0.532 ± 0.183 26. 5PW / 5PL 0.456 - 0.695 0.524 ± 0.062 0.467 - 0.612 0.539 ± 0.033 27. 3PML / ICW 0.332 - 0.435 0.392 ± 0.034 0.313 - 0.435 0.389 ± 0.036 6.3.2. Scylla olivacea (Herbst, 1796) description by Keenan et al. (1998)

Keenan et al. (1998) described S. olivacea as "Frontal lobe spines low (mean height approximately 0.03 times frontal width measured between medial orbital sutures), rounded with shallow interspaces. Antero — lateral carapace spines broad, with outer margin convex. Carpus of chelipeds usually with one small blunt prominence (may be spinous in juveniles) ventro — medially on outer margin; reduced second spine may be present dorso — distally in juveniles and young adults. Palm of cheliped usually with a pair of blunt prominences on dorsal margin behind insertion of the dactyl, inner larger than outer; may be spinous in juveniles and young adults.

Chelipeds, legs and abdomen all without obvious polygonal patterning for both sexes.

Colour varies from red through brown to browny / black depending on habitat." In addition to morphological characters, Keenan et al. (1998) used three morphometric ratios namely ICS / OCS, FMSH / FW and FW / ICW to distinguish S. olivacea from its congeners. The ratios assigned to S. olivacea are 0.006 ± 0.035, 0.029 ± 0.005 and

0.415 ± 0.017, respectively.

Joel and Raj (1980) observed chromatophores immediately below the tip of G1 of S. serrata thereby rendering brownish red colouration. However, Keenan et al.

(1998) re — identified S. serrata of Joel and Raj (1980) as S. olivacea.

6.3.3. Morph 2

6.3.3a. Material examined

One male (CW 4.13 cm), 8 May 2005, Mormugao Port Trust, Goa; one female (CW

10.46 cm), 25 February 2006, Potential fishing ground off Goa; one female (CW

10.84 cm), 27 March 2006, Potential fishing ground off Goa; one male (CW 6.43 cm), one female (CW 5.94 cm), 26 September 2006, Potential fishing ground off Goa; four

105 males (CW 9.87 cm, CW 8.63 cm, CW 10.94 cm, CW 8.79 cm), 16 January 2008, Panaji market, Goa; seven males (CW 10.26 cm, CW 10.09 cm, CW 10.61 cm, CW 10.85 cm, CW 9.72 cm, CW 9.20 cm, CW 9.10 cm), four females (CW 11.15 cm, CW 10.39 cm, CW 10.32 cm, CW 11.17 cm), 15 March 2008, Panaji market, Goa.

6.3.3b. Description

Carapace broader than long, with prominent H — shaped groove present on the cardiac region (Fig. 6.6a,b). Frontal margin of carapace (excluding inner supra — orbital angles) with four rounded spines separated by rounded interspaces, their height approximately 0.026 — 0.066 times frontal width measured between orbital sutures (Fig. 6.7). Antero — lateral margins of carapace longer than the postero — lateral margins, and divided into nine sharp spines; all spines almost equal in size. Chelipeds massive, smooth, longer than legs; merus with three spines on its anterior margin and two on the posterior margin; carpus with acute spine at the inner angle, its outer angle either lacks or may possess rudimentary spines (Fig. 6.8); propodus with a strong spine at the articulation with carpus, two reduced spines may be present on the dorsal margin behind insertion of the dactylus (Fig. 6.8). Four pairs of pereiopods, first three pairs ambulatory and similar and the fourth pair natatorial. Male abdomen bluntly triangular in shape; female abdomen watch — glass shaped (Fig. 6.9). G1 with long and slender distal portion comparatively narrower than in S. serrata, ends in long, • bluntly pointed tip; outer margins of apex convex (Fig. 6.10). Chromatophores present immediately below the tip of G1 and render a brownish red appearance in fresh specimens (The brownish red colouration fades in preserved specimens). Colouration of carapace varied, greenish brown to dark brown and generally devoid of polygonal patterning (Fig. 6.6a,b). Some specimens display light yellow

106 (i) and (ii) Absence of polygonal markings on chelipeds and legs

Fig. 6.6. Scylla olivacea (a) Dorsal view of carapace (Photograph), (b) Frontal view of carapace (Photograph)

(i) Rounded interspaces (ii) Rounded frontal spines

Fig. 6.7. Scylla olivacea — Frontal margin of carapace (Camera lucida diagram)

(ii)

(i) Blunt propodal spines behind dactyl insertion Absence of polygonal pattern (ii) Absence of spines on outer margin of carpus on female abdomen

Fig. 6.8. Scylla olivacea Fig. 6.9. Scylla olivacea — Colour

— Right cheliped (Photograph) pattern on female abdomen (Photograph)

1 mm

5 mm

(b) (c)

(i) Long, slender distal portion of G1 (ii) Long and bluntly pointed apex (iii) Slightly convex outer margin

Fig. 6.10. Scylla olivacea (a) Entire Gl, (b) Tip of Gl, (c) Tip of G1 (enlarged view) (Camera lucida diagrams) polygonal marking on the epibranchial region of carapace and the chelipeds. Ventral surface of male abdomen cream coloured, whereas, in females, the abdomen is characterized by alternating transverse dark and light green or brownish bands, and lack of polygonal patterning (Fig. 6.9). Chelipeds brown in colour and generally devoid of pOlygonal patterning (Fig. 6.6a,b) however some specimens display indistinct patterning. Legs brown and devoid of polygonal patterning (Fig. 6.6a,b). Details of morphometric measurements (range, mean and standard deviation) of the new variety of S. olivacea are provided in Table 6.3. Based on the above description, it is evident that the present specimens resemble S. olivacea's description by Keenan et al. (1998) in "lacking two well developed spines on distal half of outer margin of carpus of cheliped", "frontal lobe spines rounded with shallow interspaces", "palm of cheliped usually with a pair of blunt prominences on dorsal margin behind insertion of the dactyl", and "chelipeds, legs and abdomen without obvious polygonal patterning".

However, they differ from S. olivacea in the following: 1. "Frontal lobe spine height approximately 0.026 — 0.066 (0.045 ± 0.012) times frontal width measured between orbital sutures (FMSH / FW), as compared to

0.018 — 0.037 (0.029 ± 0.005) in S. olivacea (Keenan et al., 1998)." 2. "May possess rudimentary inner carpus spine or tubercle (range 0.000 — 0.500;

0.101 ± 0.177), whereas S. olivacea (Keenan et al., 1998) lacks any (range 0.000 — 0.250; 0.006 ± 0.035).

3. "Chromatophores just below tip of the first pair of abdominal appendages of male give a brownish red appearance in fresh specimens (faded in preserved specimens)."

107 6.3.4. Statistical analysis

The statistical analysis (Student's t — test) carried out to distinguish between the two Scylla species based on twenty seven morphometric ratios (Table 6.3), revealed that the two species differed significantly with respect to four morphometric ratios namely FMSH / FW, ICS / OCS (P < 0.001), FMSH / DFMS and OPS / PL (P

6.3.5. DNA sequencing

In view of the minor morphological differences between the present specimens of S. olivacea and the description of the same provided by Keenan et al. (1998), 18S rDNA sequencing was carried out. The DNA sequence is 385 bases long (Fig. 6.11a).

The BLAST search for the above sequence at the NCBI website revealed that S. olivacea (NCBI Accession No. AF109321) was its closest homologue with sequence match score of 0.92 (Fig. 6.11a,b).

6.3.6. Geographical distribution of Scylla olivacea

Scylla olivacea's known geographical distribution (Keenan et al., 1998) includes coasts of Pakistan, Thailand, Singapore, Malaysia, Vietnam, southern China,

Taiwan, Philippines and western Australia (Fig. 6.12). Joel and Raj (1980) reported two species of Scylla namely S. serrata and S. tranquebarica from the Pulicat lake, southeast coast of India. Keenan et al. (1998) re — identified S. serrata specimens from Joel and Raj (1980) as S. olivacea. Therefore, the present report of S. olivacea along Goa coast is the first record for the entire west coast of India.

108

ALIGNED SEQUENCE DATA: (3S5 bp) PHYLOGENETIC TREE

T:CT:.7.71a:A.SCT;;C:T.CA:27A7AP-7.GACATT:GTAkr:CAACA:CGAGfi:Cr.CAALCTCCT.:CTriclakkG FACT: 71)%75..ASFIATir:F.C: riC17,a7TA:C2C1704.:;:kkCST GATCTT:T.A.h.a... .11.77M5Cr...:C27171lakit • (Halides blinded. :GG7.GCCCCAACCGA.;:la 77AT7CGW-U.F.7:G:;.717.ATCC.ACSCII.F.TC72,72...V.GC7.717.TAGG.SG.:1").G A7CGaZ7:C:ATZ:TA:C =ITT CA: TIF.C.GLitrg:GTATT CTS:C4C4T7CGC-G • Charybdis helledi .. a GGV:TAG.G.SCGCGT:

• Postrinus trituberoulatus ALIGNMENT VIEW and DISTANCE MATRIX TABLE: • (With Sample GS2 sequence taken o reference 7.mmence) • Poltinus sayl 4 Sig. scare Organism Name NCBI Accession No • Callinactos ornetus

0.92 Scylla oltvacea AFI09321 • Callinactas maracaltroensis 0.85 Scylla nanquebarica )FI09320 0.84 Scylla paramatuo:.ain AY841366 0,81 Scylla sonata A109318 • Scylla senate 0.30 Chall.iXii5 hellsi DQ307666 0.79 Pommus tritubemilanz AY303612 Scrila pararnemosaln 0.79 Pommus lay, DQ3S8053

0.78 Callittecte °mann. DQ407679 • Scylla tranqueberica 0.85 Ovalipes punctanr, DQ062733 • 0.78 Callinectes maracaiboensis emblAT2931s3.1pAa2 98 I 85 • Whiff I 4

(a) • Scylla alivacse (b)

Fig. 6.11. DNA sequence of Scylla olivacea obtained during the present study (a) Aligned sequence data (385 bp), Alignment view and Distance matrix table, (b) Phylogenetic tree indicating position of the present sample

GOA

Fig. 6.12. Map indicating the worldwide distribution of Scylla olivacea 6.4. Conclusion

The present observations indicate a new record of S. olivacea from the entire west coast of India and further emphasize that the mud crabs from the present study area (Goa) are represented by two species, S. serrata and S. olivacea.

109 Chapter 7. Puffer fish diversity off Goa 7.1. Introduction

Puffers are a unique group of fishes those inhabit freshwater, marine and estuarine environments in tropical and temperate regions (Nelson, 1994). These are small to moderate — sized fishes, with a heavy blunt body, rounded in cross section with large, broad and blunt head (Talwar and Jhingran, 1991). These fishes are capable of rapid inflation by intake of water or air (Nelson, 1994; Shipp, 2003).

Recently, some of these fishes have acquired importance owing to nuisance created by their voracious preying habits and net — damaging capabilities (Naik, 1998).

7.2. Literature review

Published literature pertaining to puffer fishes from the Indian region revealed comprehensive taxonomic works during the nineteenth century (Russell, 1803;

Hamilton, 1822; Bleeker, 1853; Blyth, 1860b; Day, 1865, 1876 — 1878, 1889).

Dekkers (1975) reviewed the taxonomy of the genus Tetraodon through exhaustive morphological descriptions of freshly collected and museum specimens along with review of existing literature, and revealed great degree of polymorphism among puffers. Recently, Talwar and Jhingran (1991) provided descriptions of eleven puffer species of the Indian coasts along with brief information on their geographical distribution and ecology.

Recently however, molecular techniques involving DNA sequencing of puffer tissue (Boeddrich et al., 1999) have been used in conjunction with traditional taxonomy to substantiate the latter. Crnogorac — Jurcevic et al. (1997) carried out sequencing and cloning of the mitochondrial cytochrome b gene of Tetraodon fluviatilis in order to evaluate the evolutionary status of puffers. Chou et al. (1998) isolated three novel genes from the genomic DNA of T. fluviatilis. Yamanbue et al.

110 (2006) determined the entire mitochondrial genome sequence for T. nigroviridis in order to estimate phylogenetic relationships and divergence times among major lineages of fishes, whereas Chang et al. (1997) determined the genomic structure and promoter region of a single gene (c fos gene) in 7'. nigroviridis.

In view of the above, the present chapter provides a detailed account of the puffer fish diversity along Goa coast using traditional taxonomic methods. Further, the ambiguous taxonomic status of Tetraodon species using traditional criteria coupled with the effectiveness of the molecular - based techniques prompted the use of DNA sequencing.

Terminology used in the morphological description of puffer fish follows

Dekkers (1975), Froese and Pauly (2010). The following abbreviations are used. TL -

Total length; FL - Fork length; SL - Standard length; PDL - Pre - dorsal length; PAL

- Pre - anal length; PPL - Pre - pectoral length; BD - Body depth; HL - Head length;

OD - Orbital diameter; POL - Pre - orbital length; PHL - Post - orbital head length;

CH - Caudal fin height; CA - Surface area of caudal fin; AR - Aspect ratio; D -

Dorsal fin; A - Anal fin; P - Pectoral fin; C - Caudal fin

7.3. Taxonomy

73.1. Family Tetraodontidae Fritzsche, 1982

The Family Tetraodontidae owes its name to the word "Tetraodon", which in

Greek means "four teeth" (Talwar and Jhingran, 1991). Fishes of this family possess characteristic jaws wherein the teeth are fused into a beak - like dental plate with a median suture on each jaw (Talwar and Jhingran, 1991), thereby giving the appearance of four heavy, powerful teeth, two in each jaw (Shipp, 2003). In addition,

111 these fishes lack pelvic fin and their body is generally covered with spiny patches on their back, sides and belly (Talwar and Jhingran, 1991; Froese and Pauly, 2010).

The Tetraodontidae is a large group comprising one hundred and eighty seven species belonging to twenty nine genera (Froese and Pauly, 2010). Among these, twenty five species belonging to nine genera are known to inhabit the Indian waters

(Froese and Pauly, 2010). In view of the diversity of puffers in the Indian waters, a taxonomic key incorporating all genera those listed in published literature (Talwar and Jhingran, 1991; Matsuura, 2001), along with minor modifications is provided below.

1. Dorsal fin with 35 to 36 rays; anal with 28 to 29 rays Xenopterus

Troschel (ex Bibron), 1856.

- Dorsal and anal fins each with 7 to 18 rays 2.

2. Nasal organ barely visible without aid of magnification; dorsal surface

posterior to eyes distinctly keeled Canthigaster Swainson, 1839.

Nasal organ easily visible without magnification; dorsal surface posterior to

eyes more or less smooth, without a distinct keel ..3.

3. Nasal organ a short rounded tube with a terminal opening Tetraodon

Linnaeus, 1758.

- Nasal organ an upright sac with two openings (nostrils), or a solid bifid

tentacle, or an upraised cup with two fleshy lobes 4

4. Nasal organ covered by small sac with two nostrils 5

- Nasal organ not covered by a small sac with two nostrils 12

112 5. A raised skin fold along lower side of caudal peduncle .6.

- No raised skin fold along lower side of caudal peduncle 7.

6. Top of pectoral — fin base above lower margin of eye Marilyna Hardy, 1982.

- Top of pectoral — fin base below lower margin of eye .8.

7. No spinules on head or body .Sphoeroides Anonymous (Lacepede), 1798.

- Spinules all over head and body except for caudal peduncle Tylerius Hardy, 1984.

8. Mouth supra — terminal ..Amblyrhynchotes Troschel (ex Bibron), 1856. - Mouth terminal ..9.

9. Chin distinct Torquigener Whitley, 1930.

- Chin indistinct 10.

10.Lower half of body silver white, contrasted to dark dorsal half...Lagocephalus

Swainson, 1839.

- Body variously coloured, but lower half of body not silvery white ..11.

11.Spinules on back and belly well separated Taktfugu Abe, 1949.

- Spinules cover entire body except for caudal peduncle... Tetractenos Hardy, 1983.

113 12. A single lateral line on side of body; nasal organ in the form of bifid tentacle,

with minute pits on the apposing surface of each tentacle...... Arothron

1841.

Two lateral lines, the upper joining the lower in the region above or behind

anal fin; nasal organ in form of a depression with a slightly raised margin

expanded before and behind, into a pair of elongate flaps, or cup - shaped with

the front and rear edges produced into broadly rounded flaps Chelonodon

Muller, 1841.

Further, species level identification aided by published literature (Talwar and

Jhingran, 1991; Froese and Pauly, 2010) revealed four species of marine puffers namely Arothron immaculatus (Bloch and Schneider, 1801), Chelonodon patoca

(Hamilton, 1822), Lagocephalus spadiceus (Richardson, 1845), Talafugu oblongus

(Bloch, 1786), and an unidentified freshwater (Tetraodon) species. A brief description of all marine species is provided below along with published information pertaining to their geographical distribution, eco - biology and commercial importance (Froese and

Pauly, 2010). In addition, details of the morphometric measurements of the specimens collected during the present study are provided in Table 7.1.

7.3.2. Arothron immaculatus (Bloch & Schneider, 1801)

Material examined: One unsexed specimen, 29.36 cm TL, 6 January 2007, Potential fishing grounds, Goa.

Fin formula: D 0 9 - 11; A 0 9 - 17; P 19; V 0

Morphological characters: Small - sized species (maximum 30.0 cm TL) with broad and heavy body, tentaculate nasal organ; inter - orbital space flat; lateral line

114 Table 7.1. Morphometric measurements of marine puffers obtained during the present study along with their means and standard deviation

Sr. Morphometric Species No. parameters (cm) Arothron Chelonodon Lagocephalus Takifugu immaculatus patoca spadiceus oblongus (N= 01)* (N = 06) (N = 05) (N = 01)* 1. Total length 29.36 12.79 ± 5.35 10.39 ± 1.51 11.44 2. Fork length 29.36 12.79 ± 5.35 9.90 ± 1.38 11.44 3. Standard length 22.66 10.35 ± 4.40 8.37 ± 1.23 8.91 4. Pre - dorsal length 15.68 7.52 ± 3.29 5.60 ± 0.79 6.34 5. Pre - anal length 18.59 7.79 ± 3.29 5.70 ± 1.00 6.20 6. Pre - pectoral 7.18 3.73 ± 1.43 2.98 ± 0.42 3.14 length 7. Body depth 7.80 3.83 ± 1.83 2.66 ± 0.50 3.04 8. Head length 6.43 3.46 ± 1.37 2.90 ± 0.37 2.81 9. Orbital diameter 1.21 0.95 ± 0.35 0.91 ± 0.14 0.71 10. Pre - orbital 2.88 1.09 ± 0.51 1.05 ± 0.20 1.09 length 11. Post - orbital 2.34 1.41 ± 0.57 0.95 ± 0.14 1.01 head length 12. Caudal fm height 10.03 3.31 ± 1.70 1.43 ± 0.48 1.71 13. Caudal fin area!' 52.24 7.79 ± 5.13 1.52 ± 0.69 3.13 14. Aspect ratio* 1.93 1.55 ± 0.58 1.28 ± 0.42 0.93

* Single specimen, hence 1.1.± S.D. not used # Surface area expressed in cm2 *not expressed in units inconspicuous; dorsal, anal and caudal fins rounded; spinules cover most of the body surface; characteristic single blotch on pectoral fin axil, and black margin on tail fin

(Fig. 7.1a).

Geographical distribution: Red sea and South Africa to Japan and Indonesia.

Ecology: Reef — associated species that inhabits mangrove — fringed estuaries, sea grass beds and coastal waters at 1 — 17 m depths; prey items include benthic invertebrates, seaweeds and detritus.

Fishery: Caught with bottom trawls, commercially exploited for aquarium trade and although toxic, it is occasionally used for human consumption.

7.3.3. Chelonodon patoca (Hamilton, 1822)

Material examined: Six unsexed specimens; 3.49 cm SL, 12 May 2005, Mormugao bay, Goa; 9.37 cm 29 December 2006, Potential fishing grounds, Goa; 7.55 cm SL,

12.45 cm SL, 14.76 cm SL, 6 January 2007, Potential fishing grounds, Goa; 14.50 cm

SL, 28 November 2008, Potential fishing grounds, Goa.

Fin formula: D 0 9 — 10; A 0 8 — 10; P 15 — 16; V 0

Morphological characters: Small — sized species (maximum 38.0 cm SL) with elongated body with broad head, evenly arched upper profile of body between snout and caudal fin; flap — like triangular nasal organ located in a depression on the snout; broad and flat inter — orbital space; two inconspicuous lateral lines on flanks, the upper one bent downward posterior to the dorsal fin and the lower one running along lower half of tail; body covered with spinules on back, throat and belly; characteristic oval — shaped yellow spots on brown back, and white belly with yellow margin (Fig.

7.1b).

115 Fig. 7.1. Marine puffer fish species obtained during the present study (a) Arothron immaculatus, (b) Chelonodon patoca, (c) Lagocephalus spadiceus, (d) Takifugu oblongus (Photographs) Geographical Distribution: East coast of Africa to Trobiand Islands and from China to northern Australia.

Ecology: Demersal species that inhabits mudflats and sandy substrates, mangrove - fringed estuaries, nearshore coastal waters, and undertakes anadromous migration; prey items include benthic copepods, barnacles, polychaetes, nematodes, gastropods, blue - green algae and seaweeds.

Fishery: Caught with bottom trawls and although toxic, commercially exploited on small scale for human consumption.

7.3.4. Lagocephalus spadiceus (Richardson, 1845)

Material examined: Five unsexed specimens; 9.02 cm TL, 9.61 cm TL, 9.35 cm TL,

21 January 2006, Potential fishing grounds, Goa; 12.47 cm TL, 11.49 cm TL, 25

February 2006, Potential fishing grounds, Goa.

Fin formula: D 11 - 12; A 11 - 12; P 17; V 0

Morphological characters: Small - sized species (maximum 19.9 cm TL) with elongate body with broad head and rounded back, low nasal papilla; two lateral lines, the upper one runs parallel to the back and the lower one forms a straight ridge connecting the chin and lower part of caudal peduncle; spinules on the back cover only half of the distance from the inter - orbital region to the dorsal fin; lunate caudal fin; characteristic greenish - olive colouration on the back, and white on the flanks and belly, fins yellowish and posterior margin of caudal fin entirely white (Fig. 7.1 c).

Geographical distribution: Red sea and South Africa to Australia; introduced from

Red sea to the eastern Mediterranean region during the early twentieth century and established there.

116 Ecology: Demersal marine shoaling species that inhabits submerged rocky patches along shallow coastal waters, occasionally enters estuaries, and undertakes oceanodromous migration; prey items include teleosts (mackerel) and benthic invertebrates.

Fishery: Caught with bottom trawls, occasionally with shore seines, and lacks commercial value; known for net — destroying menace (Naik, 1998).

7.3.5. Takifugu oblongus (Bloch, 1786)

Material examined: One unsexed specimen, 11.44 cm TL, 19 February 2007, Aguada bay, Goa.

Fin formula: D 12 —13; A 10 —11; P 16; V 0

Morphological characters: Small — sized species (maximum 40.0 cm TL) with elongate body, broad head, slightly convex to flat inter — orbital space; spine — covered patches on the top of head, anterior part of back and belly; characteristic white spots or elongated patches on brownish back, yellowish white belly, orange coloured fins (Fig. 7.1d).

Geographically distribution: South Africa to Japan and Australia.

Ecology: Demersal marine species that inhabits shallow coastal waters, and enters estuaries; prey items include bivalves, gastropods, polychaetes, teleosts and seaweeds.

Fishery: Caught with bottom trawls and lacks commercial value.

In addition to the above, a freshwater species (N = 04) commonly referred to as "green spotted puffer" was obtained from a gill net operation at Betim

(15°30'18"N, 73°49'52"E) in December 2005, and trawl samplings (N = 02) off

Mormugao Port Trust in September 2006 and along the upper Mandovi estuary.

117 Initially, these were identified as genus Tetraodon. The identification of these puffers was carried out following the criteria suggested by Dekkers (1975) such as form of nasal organ, form of inter — orbital space, morphometric measurements, colour pattern and fin ray counts. In addition, the vertebral counts of the fish were obtained from examination of X — ray photographs of the specimens. Details of morphometric measurements of specimens along with their means and standard deviation are provided in Table 7.2.

7.3.6. Genus Tetraodon Linnaeus, 1758

Tetraodon Linnaeus, 1758 (type species: Tetrodon lineatus Linnaeus, 1758)

Fishes belonging to genus Tetraodon form a large homologous group comprising twenty three species those primarily inhabit freshwater habitats throughout the Indo — Pacific region (Dekkers, 1975; Froese and Pauly, 2010).

Dekkers (1975) defined the genus Tetraodon as: "A genus of the family Tetraodontidae with the sphenotic expanded laterally beyond frontal to form a broad flattened lobe; orbital roof scarcely arched, the lateral ethmoid not bent down before the eye, not forming an olfactory foramen; mesethmoid broad; upper lateral line not reaching end of tail, curved down above anal fin to meet lower line, which extend forward beyond the point of union; 19 vertebrae; dermal ossifications of back bearing simple prickles, evenly scattered; spines on sides small, closely set; dorsal fin maximally counting 16 rays; nasal organ an elevated tube, terminally often divided into two lips or lobes which may nearly reach the basis." He described twelve species and one sub — species and categorized them into five broad "species — groups" based on differences in morphological characters. A modification of the taxonomic key to

118 Table 7.2. Morphometric measurements of Tetraodonfluviatilis fluviatilis obtained during the present study (N = 04) along with their means and standard deviation

Sr. Morphometric ZMPT -1 MBT -1 MBT - 2 MTP Ft 1 S.D. No. parameters (cm) 1. Total length 12.55 12.38 13.51 14.40 13.21 ± 0.94 (= Fork length) 2. Standard length 9.70 9.28 10.53 10.95 10.12 ± 0.76 3. Pre - dorsal 7.34 7.38 7.52 8.23 7.62 ± 0.42 length 4. Pre - anal length 7.19 6.93 7.75 8.14 7.50 ± 0.55 5. Pre - pectoral 3.65 3.81 4.12 4.47 4.01 1 0.36 length 6. Body depth 5.12 4.63 4.18 4.58 4.63 ± 0.39 7. Head length 3.11 3.46 3.51 3.91 3.50 ± 0.33 8. Orbital diameter 0.74 0.74 0.99 0.86 0.83 ± 0.12 9. Pre - orbital 0.91 0.83 0.84 N.A. 0.86 ± 0.04° length 10. Post - orbital 1.45 1.39 1.47 N.A. 1.44 ± 0.04° head length 11. Caudal fm height 4.55 4.52 4.69 N.A. 4.59 ± 0.09° 12. Caudal fin area# 11.27 10.67 9.64 N.A. 10.53 ± 0.82° 13. Aspect ratio* 1.84 1.91 2.81 N.A. 2.19 ± 0.54° # Surface area expressed in cm2 *not expressed in units °N = 03 the five "species groups" of the genus Tetraodon (Dekkers, 1975) is provided below.

1. Nasal organ a short, rounded tube with a terminal opening, not or only slightly

two - lipped ..cutcutia - group.

- Nasal organ a short or oblong, slightly or strongly compressed tube, the distal

1/3 to 5/6 of which is divided into two lips or lobes 2.

2. Nasal tube divided over 1/3 to 1/2 of its length; apposed surfaces of nasal

lobes never with spongy tissue; sides with many polygonal dark spots leaving

only lighter network or wide - meshed dark network enclosing some dark

rounded spots; head deflated leiurus - group.

Nasal tube divided over 1/3 i. e. 1/2 to 4/5 of its length; apposed surfaces of

nasal lobes sometimes with spongy tissue; sides coloured otherwise; head

mostly rounded 3.

3. Sides with a single or double longitudinal band separating a uniformly dark

dorsal region from a differently coloured ventral region; D 9 - 11; A 8 - 9; P

18 -19 erythrotaenia - group.

Sides sometimes with a simple longitudinal band, but otherwise differently

coloured; D 11 - 16; A 10 -14, P 16 - 24 4.

4. Sides with about 35 dark transverse bandlets on upper parts; back dark; D 12;

A 11; P 16 waandersii - group.

- Sides and back with dark spots and / or blotches; D 11 - 16; A 10 -14; P 18 -

24 fluviatilis - group.

119 The above mentioned key revealed that the present specimens belonged to the fluviatilis — group. A modified version of the key to species of the fluviatilis — group

(Dekkers, 1975) is provided below.

1. Sides with two to four dark ocellated spots and with irregular whitish lines, some

of them broken into spots; back with similar lines and with three dark patches; D

13-15;A 11-13;P 18-21 T steindachneri Dekkers, 1975.

Sides without whitish markings; back with or without whitish markings and dark

patches; D 1l-16;A10-14;P 18 — 24 2.

2. Back and sides with more or less regular, rounded spots, which are more or less

evenly scattered, and only occasionally confluent, never forming broad blotches;

apposed surfaces of nasal lobes often spongy; D 12 — 14; A 10 — 12; P 18

23 T. nigroviridis Marion de Proce, 1822.

Back with dark transverse markings or with large dark blotches; nasal lobes with

or without spongy tissue; D 11 — 16; A 10 —14; P 18 — 24 3

3. Back with dark transverse bands, often including a V — shaped marking behind

eyes; apposed surfaces of nasal lobes always without spongy tissue in all

specimens 60 cm SL; often a dark broad longitudinal band separating dark

colour of upper parts from light colour of belly; D 11 — 13; A 10; P 18 — 19...... T.

kretamensis Inger, 1953.

Back with three to four dark blotches in longitudinal row; apposed surfaces of

nasal lobes with spongy tissue in 85 % of specimens _?:60 cm SL; never a dark

longitudinal band; D 13 — 16; A 11 — 14; P 19 — 24 T. fluviatilis Hamilton,

1822.

120 Further, within the fluviatilis - group, the present specimens resembled T. fluviatilis based on colour pattern and fin ray counts. Dekkers (1975) reported two sub

- species of T. fluviatilis namely T. fluviatilis fluviatilis Hamilton, 1822 and T. fluviatilis sabahensis Dekkers, 1975. The present specimens showed greatest resemblance to Dekkers' description of T. fluviatilis fluviatilis.

7.3.7. Tetraodon fluviatilis fluviatilis Hamilton, 1822

Z3. 7a. Dekkers' (1975) description

Dekkers (1975) described T. fluviatilis fluviatilis as "Body oblong, compressed laterally except in fleshy specimens; dorsal profile arched, highest at midst of back; inter - orbital convex, without a groove; lateral line system distinct; body spines small, sometimes very distinctly two based, sometimes partially or wholly indistinct, covering back, sides and belly between eyes and origin of dorsal fin; origin of anal fin situated beneath anterior half of dorsal fin base; mouth terminal, directed forwards; lower border of eye slightly above or below level of mouth corner, upper border not interfering with dorsal profile; nasal organ a tentacle, more than distal half of which is divided in two flattened and broadened lobes; apposed surface of lobes generally with spongy tissue (in 90 % of specimens mm sl, N = 20; in

80 % of specimens < 60 mm sl, N = 5)."

"Colour in alcohol: Ground colour of upper parts tan, of belly yellowish white; three large yellowish - encircled dark patches on back between eyes and dorsal fin; anterior patch on posterior half of head (length along midline of back 6.2 - 14.0 in sl, and 1.3 - 2.9 in transverse length), middle patch between pectoral fins (length along midline of back 5.5 - 13.8 in sl, and 0.8 - 1.8 in transverse length), posterior patch in front of dorsal fin (length along midline of back 8.3 - 14.4 in sl, and 0.6 - 1.3 in

121 transverse length); middle dorsal patch occasionally broken up into two smaller patches; anterior half of head variably coloured, often irregularly dark; sides with 5 —

28 rounded, mostly ocellated, dark spots (horizontal diameter 8 — 80 in sl) of which always one at base of dorsal fin (usually the largest), one at base of caudal fin (usually the smallest), and one below or against posterior dorsal patch; belly uniform yellowish white or with same or confluent spots as on sides; 0 — 9 (m 3.8) crossbars on caudal fin; sometimes a small dark spot on interior side of pectoral fin base; other fins plain" (Fig. 7.2a,b).

7.3.7b. Present specimens — comparison with Dekkers' description

The present specimens resembled Dekkers' (1975) description of T. fluviatilis fluviatilis by virtue of the following morphological characters.

1. Nasal organ a tentacle, more than distal half of which is divided into two

flattened and broadened lobes; apposed surfaces of lobes generally with

spongy tissue.

2. Snout length almost 2.3 — 2.9 times in HL.

3. HL > 2.4 times in SL.

4. Inter — orbital region with convex profile, and lacks a groove on its surface.

5. Mouth terminal in position, directed forwards.

6. Three large dark transverse bands on back encircled by yellow colouration

(fades in preserved specimens); middle band between pectoral fins, posterior

band in front of dorsal fin (Figs. 7.3a,b, 7.4a,b).

7. Fin count D 13, A 11, P 21 — 22 (i. e. within the range described by Dekkers

(1975)).

122 (Lateral view) (a)

(iv)

(1) 3 dark transverse bands on back (II) 4 dark transverse bands on caudal fin (111) 3 dark transverse bands on back (iv) Anterior band on posterior part of head (v) Middle band between the pectoral fins (vi) posterior band in front of dorsal fin

Fig. 7.2. Diagrammatic representation of colour pattern on (a) sides and (b) back of Tetraodon fluviatilis fluviatilis (redrawn and modified from Dekkers, 1975) (a)

(i )

(b)

(i) 3 dark transverse bands on back (ii) Single broad transverse band on caudal fin

Fig. 7.3. Colour pattern on the sides of Tetraodonfluviatilis fluviatilis collected during the present study (a) Photograph (b) Diagrammatic representation (a)

(i)

(b)

(i) Anterior band between the eyes (ii) Middle band between the pectoral fins (iii) Posterior band in front of dorsal fins

Fig. 7.4. Colour pattern on the back of Tetraodon fluviatilis fluviatilis collected during the present study (a) Photograph (b) Diagrammatic representation However, they differed subtly from T. fluviatilis fluviatilis in possessing:

1. Anterior dorsal band between the eyes (Figs. 7.3a,b, 7.4a,b), as compared to

"posterior portion of head" (behind the eyes) in T. fluviatilis fluviatilis (Fig.

7.2a,b),

2. More than thirty ocellated dark patches on sides (Fig. 7.3a,b), as compared to

"5 — 28 ocellated patches" in T. fluviatilis fluviatilis (Fig. 7.2a),

3. One dark broad transverse band on caudal fin (Fig. 7.3a,b), as compared to

four bands in T. fluviatilis fluviatilis (Fig. 7.2a).

7.3.7c. DNA sequencing

The DNA sequence of Tetraodon species is 584 bases long. The BLAST search of the above sequence at the NCBI revealed that it matched with T. nigroviridis (NCBI Accession No. AP006046; Yamanoue et al., 2006; Fig. 7.5a,b).

7.4. Discussion

The present findings suggested that the nearshore waters serve as potential spawning grounds of the marine puffer L. spadiceus as evident from the capture of large shoals of adults during November — February during each sampling year. Naik

(1998) emphasized the importance of submerged rocky patches off the adjacent

Konkatt coast as spawning areas for L. spadiceus. Further, the capture of C. patoca and T. oblongus juveniles from the turbid waters of the Mandovi — Zuari estuaries suggests their importance as nursery areas. Nagelkerken et al. (2008) emphasized the importance of mangrove — fringed turbid estuarine waters as protected refuge for juveniles of diverse marine fishes owing to availability of adequate food and reduced chances of predation.

123 ALIGNED SEQUENCE DATA:MAO PHYLOGENETIC TREE

AN:ATCOOCITTTOCAMAX:AATZAATA. CCGCCTOCCCTOTC.A.M.ACCITTAAt1===T • Pampers poedloptenn AlTrIVACCGIVCAAAGC:14CCOATCACTIVIKTITAAATGOCCACCTZTAMAATCX•CtArtila=C' TIAACM,T. CCTrIlICAJattAXIVAAXIII,ATC11TCCOTOGACAACCOMAATAAWCATWACCAC PJ4ACIXI11 MACCITtAGACAWJACLCATCACCIVWCA=t1MI11TAAAW...4AtA4ACTMOt.A4 RICITITCAAATVKIT114111=ACCOM=2MACAMMAOOCCCAMTGAATAAWC.ACCCrt • Cholonorlon palm ITI.A.AMCVAGAGCCAOMCIVTMTGANZACAACATCIGACCAA MAGA =VA-CW.4=A TCAACGM COCMMTVVV. CTVAGATWAGte•C4ATI.... ■ 4 t.K.GO,CATAMGACkled444177ACCACCMCAT • Prothro,' finnementren CTMCC•11, ∎V ,, ATCCTAXIVOINX.A=P:CTATTMOCGIICCTITGTCAP.CGAITAAAGTCCTACC TrA

• Centhigester rinilato A LIGNMENT VIEW and DISTANCE AlAIRIX TABLE: (WO ( ;MAC 1 ...looter !at"( at reference sequence! • • Sphoemides macula. S.,mtv atom Orromism Nome NCItI AomOof• • • Toreador, eigrovredis 100 MM.:1M Mgt. yid is 6P0060.16 099 Tetrandon nigern irides m0(1(296015 2 0.99 Tettscdon nisrmiridir, evert( CR700241.2 • Torrnorion nigroviridis 090 l'etranden nigros nit's engs ( 1(606261 2 0.96 Arnthron firmanienrunt 11061)0742 090 Terrowl0n no Warr AV670660 1! • Tetreodon nigroviddis 0.93 erenthigager rivulattet 600,26744 094 Srboartudes maculain 1112.391114 • Tetreodon nigroviddis 0,91 I:below-4m men, 611194242 0,90 Pnrajulis ruceilerrerrn 11192032 Sob Sequence Mott( Serve (a) (b) • Totraodon euvrehlis

Fig. 7.5. DNA sequence of Tetraodon nigroviridis obtained during the present study (a) Aligned sequence data (584 bp), Alignment view and Distance matrix table, (b) Phylogenetic tree indicating position of the present sample The present results of fish identification based on the DNA sequencing were found to differ from those based on morphological observation of the specimens. Although Dekkers (1975) laid down well — defined criteria for identification of freshwater puffer fishes belonging to the fluviatilis — group (that includes both T. fluviatilis and T. nigroviridis), there exists uncertainty with regards to compliance between traditional methods and nucleotide sequencing (Crnogorac — Jurcevic et al.,

1997; Chou et al., 1998; Boeddrich et al., 1999). Moreover, the confusion prevailed as Chang et al. (1997) synonymized T. nigroviridis with T. fluviatilis. It is apparent from the observations made in the present study that there exist few differences in the morphological characters associated with the identification of closely related puffer fish belonging to T. fluviatilis group. Further, our observations also reveal that the taxonomic keys available in literature specific to T. fluviatilis group exhibit variability among the region.

7.5. Conclusion

The present observations revealed four marine and one freshwater species of puffer fishes from the coastal waters of Goa. Among these species, the present specimens of Tetraodon were found to resemble T. fluviatilis fluviatilis. The above findings verified through 18S rDNA sequencing revealed the species to be T. nigroviridis.

124 Chapter 8. Summary The present study provides authentic information on the species diversity of demersal fish of the estuarine and shelf regions (up to 25 m depth) of Goa, central west coast of India.

1. The present observations revealed altogether two hundred and four taxa

belonging to five phyla, ten classes, two clades, twenty seven orders, ninety

nine families, and one hundred and fifty nine genera, and among these two

hundred and one taxa were identified up to the species level. The highlight of

the study was fifty five new records of species to this region, among which

one was new to science and four others were first records for the entire west

coast of India.

2. The size class analysis of demersal fauna revealed the preponderance of

juveniles and adults of most invertebrates and resident fin fish, whereas among

the migrant fin fish population, juveniles and young recruits dominated, thus

suggesting the importance of these coastal waters as nursery and foraging

grounds.

3. The quantitative analysis of the trawl catches revealed the dominance of

crustaceans. However, the reduced catches of penaeid prawns indicate

reduction in stocks owing to intensive exploitation since the introduction of

mechanized vessels. Further, the dominance of other groups (fin fish, other

crustaceans, molluscs and other fauna) in the trawl catches during the present

study elucidates the non — selective nature of the fishing gear.

4. The present observations revealed that the demersal faunal community of Goa

coast is characterized by high species richness (2.04 ± 0.66), diversity (1.94 ±

0.44) and low evenness (0.05 ± 0.02).

125 5. The present computed fish catch rate of 99.47 kg.hf l indicates a sharp decline of 51.27 % during the past three decades, primarily attributed to a significant increase in the numbers of mechanized fishing vessels. Further, reduction in catches of higher carnivores coupled with phenomenal increase in lower trophic level catches reflects gradual replacement by planktivores. 6., The taxonomic diagnosis of rarely occurring taxa revealed a portunid crab

species namely Charybdis goaensis — a species new to science. The comparison of its morphological description with forty two extant congeners

revealed closest resemblance to C. philippinensis, which was re — described to facilitate thorough comparison with the new species.

7. A rare reef — inhabiting fusilier namely Caesio cuning was recorded for the first time from the Goa coast and outside its known geographical array. The occurrence of reef fish along the Goa coast was attributed to the artificial reef

— like substratum created by the grounded vessel MV River Princess.

8. A new record of the mud crab Scylla olivacea was found from the entire west coast of India; however minor differences in morphological characters prompted the use of 18S rDNA sequencing. The latter technique corroborated the results of the orthodox traditional taxonomic method. 9. The puffer fish diversity along Goa coast revealed four marine species namely

Arothron immaculatus, Chelonodon patoca, Lagocephalus spadiceus,

Takifugu oblongus along with an unidentified Tetraodon species. Comparison

of present Tetraodon specimens revealed close resemblance to T. fluviatilis

fluviatilis. The above findings verified through 18S rDNA sequencing

revealed the species to be T. nigroviridis.

126 Bibliography Achuthankutty C.T. and Nair S.R.S. (1993) A new record of two penaeid species from Goa coast. Journal of Indian Fisheries Association, 23: 109 —111.

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158 Published articles Padate V.P., Baragi L.V. and Rivonker C.U. (2009) Analysis of few biological aspects of sea snakes caught incidentally by commercial trawlers off Goa, West coast of India. Journal of Threatened Taxa, 12 (1): 606 - 612. Padate V.P., Rivonker C.U. and Anil A.C. (2010a) A note on the occurrence of reef inhabiting, red — bellied yellow tail fusilier, Caesio tuning from outside known geographical array. Marine Biodiversity Records, e3 (21): 1— 6. Padate V.P., Rivonker C.U., Anil A.C., Sawant S.S. and Venkat K. (2010b) A new species of portunid crab of the .genus Charybdis (De Haan, 1833) (Crustacea: Decapoda: Brachyura) from Goa, India. Marine Biology Research, 6 (6): 579 — 590.

Oral and Poster Presentations Padate V.P., Rivonker C.U., Anil A.C., Sawant S.S. and Venkat K. (2008) Spatio — temporal variations in demersal fish community along the fishing grounds of Goa, West coast of India. Oral Presentation at the International Conference on Biofouling and Ballast Water Management held at the National Institute of Oceanography, Dona Paula, Goa during 5 — 7 February, 2008. Padate V.P. and Rivonker C.U. (2008) Demersal Marine Fish Community from Goa, Central West Coast of India — A search for Rare Species. Poster Presentation at the International Conference on Biofouling and Ballast Water Management held at the National Institute of Oceanography, Dona Paula, Goa during 5 — 7 February, 2008. Rivonker C.U. and Padate V.P. (2009) Demersal marine fauna off Goa coast. Poster Presentation at the National Workshop on Ballast Water Management held at the National Institute of Oceanography, Dona Paula, Goa on 16 November, 2009. Rivonker C.U. and Padate V.P. (2009) Inventory of new records of demersal marine fauna off Goa coast. Poster Presentation at the National Workshop on Ballast Water Management held at the National Institute of Oceanography, Dona Paula, Goa on 16 November, 2009. Rivonker C.U. and Padate V.P. (2009) Demersal marine fauna off Visakhapatnam coast. Poster Presentation at the National Workshop on Ballast Water Management held at the National Institute of Oceanography, Dona Paula, Goa on 16 November, 2009. Rivonker C.U. and Padate V.P. (2009) Inventory of new records of demersal marine fauna off Visakhapatnam coast. Poster Presentation at the National Workshop on Ballast Water Management held at the National Institute of Oceanography, Dona Paula, Goa on 16 November, 2009. Padate V.P., Rivonker C.U., Anil A.C., Sawant S.S. and Venkat K. (2009) Outputs of Ballast Water Management Program, India — Published and accepted papers. Poster Presentation at the National Workshop on Ballast Water Management held at the National Institute of Oceanography, Dona Paula, Goa on 16 November, 2009. Padate V.P., Rivonker C.U., Anil A.C., Sawant S.S. and Venkat K. (2009) Outputs of Ballast Water Management Program, India — Communicated to Scientific Journals. Poster Presentation at the National Workshop on Ballast Water Management held at the National Institute of Oceanography, Dona Paula, Goa on 16 November, 2009. JoTT COMMUNICATION 1(12): 609-616

Biological aspects of sea snakes caught incidentally by commercial trawlers off Goa, west coast of India

Vinay P. Padate', Lalita V. Baragi 2 & Chandrashekher U. Rivonker 3

'.3 Department of Marine Sciences, Goa University, Taleigao Plateau, Goa 403206, India 2 National Institute of Oceanography (CSIR), Dona Paula, Goa 403004, India Email: ' [email protected] ; 2 [email protected] ; 3 [email protected] (corresponding author)

Date of publication (online): 26 December 2009 Abstract: Sea snakes occur in trawl hauls as by-catch, incurring mortality in populations inhabiting Date of publication (print): 26 December 2009 commercial fishing grounds (< 20 m depth) along the coastal inshore waters of Goa. Observations ISSN 0974-7907 (online) 1 0974-7893 (print) of this incidental catch show that true sea snakes inhabiting inshore waters comprise two species: Enhydrina schistose and Editor: Ted J. Wassenberg Lapemis curtus, contributing 65 and 35 % of the population respectively. 70 trawl operations over a period of 17 months with a total fishing effort of 110 Manuscript details: hours yielded 43 individuals, all females, which numerically contributed - 1 % to the total trawl Ms # 02253 catch. Seasonal variations indicate that there is an increasing trend in abundance from post- Received 07 July 2009 monsoon to pre-monsoon season. The capture of a gravid female from the estuarine inshore Final received 25 November 2009 waters during January suggests parturition and recruitment among sea snakes during the post- Finally accepted 27 November 2009 monsoon season. An assessment of the food composition in the stomach content revealed Citation: Padate, V.P., L.V. Baragi & C.U. completely digested prey in smaller individuals, whereas in larger-sized Individuals these items Rivonker (2009). Biological aspects of sea snakes were either undigested or semi-digested. Qualitative assessment of gut content of sea snakes caught incidentally by commercial trawlers off revealed the dominance of finfish (Ariidae, Engraulidae, Clupeidae). A biological assessment of Goa, west coast of India. Journal of Threatened a gravid female and the association of sea snakes with a barnacle species (Octolasmis grayil) Taxa 1(12): 609-616. are described.

Copyright: Vinay P. Padate , Lalita V. Baragi & Keywords: Sea snakes, Goa, trawl, seasonal variations, biological aspects. Chandrashekher U. Rivonker 2009. Creative Com- mons Attribution 3.0 Unported License. JoTT al- lows unrestricted use of this article In any me- dium for non-profit purposes, reproduction and INTRODUCTION distribution by providing adequate credit to the authors and the source of publication. Coastal ecosystems exhibit high variability in ecological parameters that are largely influenced by land use patterns and runoff, resulting in seasonal anomalies. The Author Details: VINAY P. PADATE obtained his MSc degree in Marine Sciences from Goa University prevalence of a wide range of abiotic factors provides an array of habitats for In 2005 and is currently working as Research opportunistic marine species that move between coastal and estuarine environments, Assistant at the Department of Marine Sciences, 1995; Goa University. He is engaged in studying thus providing enormous scope for increased diversity (Ansari et al. Venkataraman taxonomy and community structure of demersal & Wafar 2005). Among these areas, commercial fishing grounds are of great concern as marine fauna along the coasts of Goa and they are subjected to indiscriminate removal of non-target species (Kumar & Deepthi Visakhapatnam, India. LALITA V. BARAOI obtained her MSc degree in Marine Sciences from Goa 2006) as by-catch, including sea snakes. Sea snakes inhabit shallow estuarine waters University in 2008 and is currently working as and coral reefs throughout the Indo-Pacific region (Burns & Heatwole 1998; Heatwole Project Assistant at the Marine Corrosion and 1999). Sixty-two species have been recorded so far, and approximately 60% of these Materials Research Division, National Institute of Oceanography, Dona Paula, Goa. She is are known to inhabit Australian territorial waters (Heatwole 1999). Indian waters are engaged in studying blo-fouling organisms, known to harbour about 25 species, with the report of 11 species from the Gulf of particularly barnacles. DR. CHANDRASHEKHER U. Mannar (Karthikeyan et al. 2008) 2004). RivoNxen obtained PhD in Marine Sciences in 1992 and two along the West coast (Lobo et al. from Goa University and has about 22 Although sea snakes are commercially exploited for various applications in the publications in referred journals. Presently Philippines, Australia, Japan, Taiwan and Thailand (Rasmussen 2001), in other coastal Associate Professor, he is working in the area of areas they are considered nuisance species. Demersal fish diversity. Sea snakes show seasonal movement between inshore and offshore waters either in Author Contribution: VPP carried out sampling search of food or for bearing young (Shuntov 1971; Wassenberg et al. 1994). Fry et al. for the study along the potential fishing grounds off Goa, followed by taxonomic Identification of (2001) suggested that the females of most species appear to be gravid in the summer, sea snakes and their seasonal distribution. LVB and presumably bear young towards the end of this period in Australian waters. carried out the biological analyses including gut Sea snakes form an important constituent in the marine environment as they occupy content analysis, reproduction and symbiotic associations. CUR planned, designed and a high position in the food web, feeding upon various finfish and invertebrate species participated In the field work and was instrumental (Glodek & Voris 1982; Heatwole 1999). In addition, sea snakes interact with other In interpretation and discussion of the manuscript. marine organisms through symbiotic interactions. For example, sea snakes are reported Acknowledgement: See end of this article. to be fouled by barnacles (Reynolds & Pickwell 1984) and other marine organisms (Denson 1975; Zann et al. 1975). The present paper addresses the taxonomic composition of the assemblage of true sea snake species, seasonal variations in trawl catches and certain biological attributes such as feeding habits, reproduction and symbiotic associations, based on observations made using commercial trawlers along the fishing grounds of Goa.

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Journal of Threatened Taxa I www.threatenedtaxa.org I December 2009 I 1(12): 609-616 609 Sea snakes of west coast of India V.P. Padate et al.

MATERIALS AND METHODS each in Aguada and Mormugao bays, three each in Mandovi Estuary and Cumbharjua Canal) during September and October Study area: The coastal waters of Goa are flanked by six 2006, and May, September and December 2007 and January estuaries, the Mandovi-Zuari estuarine complex (15°25'- 2008. Trawl nets with mesh sizes of 15mm (mouth end) and 15°31'N & 73°45'-73°59t, Fig. 1) being the most prominent 9mm (cod end) were towed at a speed of approximately 2 knots ecosystem in terms of productivity and species diversity, (4 km Ill. The catch obtained was sorted into five sub-samples. possibly due to regular tidal inundation (pers. obs.) in spite of Uncommon (or rare) specimens including sea snakes were the influence of anthropogenic input. During the South-west picked out, put on ice and sent to the laboratory for detailed monsoon, fresh water influx becomes a major factor that controls examination. In addition, samples were obtained by operating the hydrodynamics of this estuarine complex, whereas tides beach seines along Betim (15°30'18"N & 73°49'52"E) during play an important role during other periods. The bathymetry December, 2005 and twice in the vicinity of Mormugao Port of this estuarine complex indicates silt, clay and detritus Trust (15°24'16"N & 75°48'56"E) during December, 2005 and transported by riverine influx from the upper reaches where September, 2006 to assess the occurrence of demersal species mangrove vegetation occurs in high density, making for a in the estuarine embayment, as these were inaccessible to highly productive detritus-based food chain (Kulkarni et al. demersal trawlers. Sampling could not be undertaken during 2003) with highly diversified biota. The study area monsoon due to a ban on trawl operations along the Goan coast encompasses the fishing grounds < 20m depth that are (Goa, Daman and Diu Marine Fishing Regulation Rules 1982). subjected to increasing levels of exploitation by commercial Voucher specimens were stored in 5% formalin solution in trawl operations. transparent plastic bottles. These are deposited at the Marine Sampling strategy: During the study period (February Biology laboratory, Department of Marine Sciences, Goa 2006 to February 2008) a total of 70 trawl operations (110 h U niversity. fishing effort) were undertaken onboard commercial trawlers Species identification: At the laboratory, specimens were in the estuarine and inshore waters of Goa ; west coast of India preserved using 6 - 8 % formalin solution and identified up to (Fig. 1) to assess the diversity and total community structure the species level by employing orthodox taxonomic methods of the demersal fauna. Twenty trawl operations were once a such as morphometric and meristic characters aided by an month (during January to April 2006, October 2006, and January identification key (Rasmussen 2001). Subsequently, snout-vent 2008) and 50 were undertaken on fortnightly basis (during length and tail length (in centimetres) of each specimen were December 2006 to May 2007, November 2007, and February measured using measuring tape and total weight (in grams) 2008). Sixteen trawling operations were undertaken in the was measured using an electronic balance (Sartorius - CP225D). Mandovi-Zuari estuarine complex (six in Zuari Estuary, two Gut content analysis and data compilation: In order to

Figure 1. Map of study area indicating trawl sampling sites 610 Journal or Threatened Taxa I www.threatenedtaxa.org I December 2009 I 1(12): 609-616 Sea snakes of west coast of India V.P. Padate et al.

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Image 1. Enhydrina schistose with distinguishing character (inset)

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Image 2. Lapemls curtus with distinguishing character (inset) analyze dietary items of sea snakes, the specimens were first Table 1. Taxonomic characters of sea snakes observed dissected to expose the gut, then the gut was divided into three along Goa coast parts (foregut, midget and hindgut) and the status of food items (undigested, semi-digested or completely digested) examined. Taxonomic character Enhydrina schlstosa LapernIs curtus Food contents were subjected to quantitative assessment using 1. Scales around neck 45 - 50 25 - 30 a wet gravimetric method (Hyslop 1980). Undigested prey 2. Scale rows around body 50 - 55 40 - 42 was identified to species level. 3. Number of ventral scales 250 - 270 200 - 220 4. Distinguishing characters Mental shield on ventral Mental shield The data collected on sea snakes and the trawl catch margin of head not elongated. composition during the study period (except monsoon) was used elongated. All scale rows of Lower scale to describe seasonal variations. These data were assessed in similar size. rows enlarged. terms of the number per trawl and later expressed as a percentage contribution. In addition, demersal fishes occurring in the trawl catch and those found in the gut of sea snakes were and their prey items composition within the catch was collated identified to species level using identification keys (Fischer & to respective seasons and plotted graphically. Whitehead 1974). The monthly data collected on sea snakes Analysis of reproduction: In order to have an insight into

Journal of Threatened Taxa 1 www.threatenedtaxa.org I December 2009 1 1(12): 609-616 611 Sea snakes of west coast of India V.P. Padate et al.

Table 2. Details of morphometric measurements of sea snake specimens (N=43) collected from Goa coast along with their means and standard deviation

Morphometrlc Species parameters E. schistose L . curtus Range Means S.D. Range Mean S.D.

1. Snout-vent length (cm) 38.75 71.25 56.34 * 9.57 42.00 - 48.75 45.70 * 2.30 2. Tall length (cm) 3.75 - 12.50 7.95* 2.89 3.75 - 7.50 5.30 s 1.05 3. Weight (g) 32.00 - 150.00 98.18 ± 37.43 40.00 - 120.92 60.05 ± 26.19 reproductive behaviour, the sex of each specimen was 50' determined based on anatomical features (hemipenes in males and ovaries in females). In the case of gravid females, uterine tubes containing ova were separated from the viscera and the diameter (cm) and weight (g) of each ovum measured using vernier callipers and an electronic balance, respectively. Symbiotic associations: Each sea snake was observed for the presence of barnacles. These were detached from skin of the specimen using forceps, and identified to species level using the identification key provided by Fernando (2006).

RESULTS AND DISCUSSION

Species composition, abundance and seasonal trends of sea snakes True sea snakes (Hydrophiidae) form an important component of the coastal habitats of tropical and sub-tropical marine environments. These marine reptiles are known to occur in large numbers in the Indo-Pacific region (Heatwole 1999; Karthikeyan et al. 2008). However, published reports (Lobo et al. 2004) indicate that the species diversity among these groups decreases progressively towards the west. In the present study, sea snake samples contained only females of two species (Table 1), namely Enhydrina schistosa (Image 1) and Lapemis curtus (Image 2). The present observations suggest that the collected individuals (all females) were mainly involved in inshore PRM POM PRM estuarine migrations. Voris et al. (1978) indicated that estuaries serve as a nursery and that adults enter estuaries to bear young. 2007------I -2008-i Lobo et al. (2004) reported the occurrence of these two species Sampling season along the Goan coast, however, no attempt was made to provide :-64,;•- Sea catfishes Sardines and Herrings Anchovies information on the systematics of these species. They further Sea snakes suggested that sex ratios of both the species were highly biased -40- Left-handed soles Swimming crabs Penaeld prawns towards females, whereas Voris & Jayne (1979) and Lemen & PRM Premonsoon POM - Postmonsoon Voris (1981) reported seasonal fluctuations in the sex ratio of sea snakes from Malaysia and suggested that this could be a Figure 2. Seasonal variations In percentage contribution of function of sexual differences in habitat selection and their sea snakes and their prey items to total trawl catch activity. The present observations were made over 17 months with estuaries and shallow coastal reefs (Rasmussen 2001) due to 110h fishing effort, and revealed only 43 individuals (all water circulation and abundance of prey (Shuntov 1971), high females), thus indicating sparse occurrence (< 1 % in terms of level of targeted fishing as well as growing incidents of their numbers per trawl) of these species in the coastal fishing incidental capture in prawn fishing grounds (Wassenberg et grounds (Table 2). Such observations also indicate that sea al. 1994) might also augment to their occurrence in low numbers. snakes are subsidiary and non-target organisms occurring in Among the sea snakes caught during the present study, E. the trawl catches (pers. obs.). Further, occurrence of these schistosa (> 65%) dominated the trawl catch as compared to L. species in low numbers in the trawl catch could be attributed to curtus (< 35%). However, Tu (1974) and Wassenberg et al. several factors such as low reproductive rate (Voris & Jayne (1994) reported that L. liordwickii (L. curtus) contributes to about 1979; Lemen & Voris 1981; Heatwole 1997), high rate of 81-88% and 53% of all the sea snakes caught in the coastal infertility (Lemen & Voris 1981), high mortality rate up to waters of Thailand and Gulf of Carpentaria, Australia, 45.8% among neonates and survival rates as low as 6% among respectively. potentially reproducing females (Voris & Jayne 1979). The numerical data collected during trawl fishing was Moreover, factors such as restricted spatial distribution in grouped under four major categories (teleosts, crustaceans,

612 Journal of Threatened Taxa I www.threatenedtaxa.org I December 2009 I 1(12): 609-616 Sea snakes of west coast of India V.P. Padate at al.

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Figure 3. Flow chart depicting hypothetical food chain and energy flow of sea snakes

Table 3. Month wise percentage of sea snakes and their dietary items in the trawl catches of Goan coast

Faunal group PRM POM PRM POM PRM 2006 2006-07 2007 2007-08 2008 F M A M A J F

Sea snakes 0.00 0.02 0.01 0.00 0.01 0.03 0.01 0.01 0.00 0.55 0.00 0.66 1.93 Sea catfishes 0.25 0.46 1.00 0.00 2.79 0.98 1.04 0.49 2.34 1.07 1.10 5.46 0.32 Anchovies 7.84 1.29 2.17 0.29 0.65 2.68 2.35 7.00 3.42 1.65 0.56 5.29 3.12 Sardines and Herrings 1.27 2.17 4.77 7.61 1.78 6.78 3.14 5.56 2.23 4.57 8.97 7.63 1.61 Croakers 0.59 1.68 2.81 1.22 4.73 5.11 6.95 2.84 1.29 2.68 2.38 4.81 2.92 Left-handed Soles 1.40 0.67 0.41 3.48 0.58 1.27 2.77 0.44 1.05 6.09 0.00 1.48 1.35 Swimming Crabs 15.04 2.94 1.30 1.61 12.61 3.92 2.97 1.96 0.77 3.19 2.07 0.84 4.25 Penaeld Prawns 22.31 64.13 54.30 27.36 32.01 31.87 28.88 42.48 53.41 29.26 31.64 13.27 31.38

PRM - Pre-monsoon (February to May); POM - Post-monsoon (November to January)

molluscs and sea snakes) indicating maximum (- 63%) along the Goan coast (pers. obs.). Another possible reason could contribution from crustaceans, followed by teleosts (- 35%), be the migration of females to the inshore estuarine waters as whereas molluscs and sea snakes contribute much less (- 1%) only female individuals were captured during the present study. each. These include a fully gravid E. schistosa female captured from The seasonal variations in the percentage of sea snakes in the estuarine waters at a depth of about 5-8 m. Ansari et al. the trawl catches reveal that the contribution of sea snakes (1995, 2003) suggest that this estuarine complex with mangrove increased from December to February (post-monsoon; Table vegetation acts as a potential nursery ground for opportunistic 3). This can be largely attributed to the commencement of marine species that migrate there either for spawning or for fishing at the end of the monsoon ban and subsequent increase feeding purpose. Voris (1985) reported feeding of E. schistosa in number of fishing boats due to calm conditions in the sea juveniles on young ariid catfish in Muar Estuary, Malaysia

Journal of Threatened Taxa I www.threatenedtaxa.org I December 2009 I 1(12): 609-616 613 Sea snakes of west coast of India V.P. Padate at al.

Table 4. Qualitative analysis of the gut contents of sea snakes collected In trawl catches of Goan coast

Sampling Species Gut content Date Foregut Midgut Hindgut

1. 27/3/2006 E. schlstosa E SD' CD 2. 27/3/2006 E. schistose E SD' CD 3. 27/3/2006 L. curtus E SD' CD 4. 24/4/2006 E. schistose E SD' CD 5. 24/4/2006 E. schlstosa E SD' CD Right 6. 6/1/2007 E. schistosa E SD' CD Left uterine uterine ' 7. 6/1/2007 L. curtus E SD' CD tube 8. 26/1/2007 E. schlstosa tube E SD' CD 9. 3/2/2007 L. curtus E SW` CD 10. 19/2/2007 E. schistose E SD' CD 11. 11/3/2007 E. schistose E SD' CD 12. 25/3/2007 L. curtus E SD' CD 13. 11/4/2007 L. curtus E SD' CD 14. 3/11/2007 E. schistose E SD' CD 15. 30/11/2007 E. schistose E SD* CD 16. 9/1/2008 E. schistose E SD' CD 17. 29/1/2008 E. schistose E SD$ CD 18. 10/2/2008 L. curtus E SD' CD

E — Empty; SD — Semi-digested; CD — Completely digested $ — Arius Jetta; — Thryssa dussumIert, * — Unidentified clupeiform fish

Image 3. Matured ova of Enhydrina schistosa along the estuarine regions only during spring and attributed this to seasonal changes in water currents and to abundance of fish.

Dietary composition and trophic level of sea snakes The qualitative analysis of gut contents of sea snakes (N = 41) revealed an empty foregut in all specimens. However, the mid guts of larger individuals contained undigested (— 25%) to semi-digested prey (— 75%), whereas those of smaller individuals contained completely digested food. In addition, the hindgut of all specimens was observed to contain completely digested food. Identification of the dietary components of sea snakes revealed two partially digested prey items namely, Arius fella (Family Ariidae) and Thryssa dussumieri (Family Engraulidae) from the mid gut of E. schistosa. Published literature on diet composition of E. schistosa (Voris et al. 1978; Voris & Voris Capitulum 1983) suggests that it prefers to prey upon catfish. Further, Octolasmis Voris et al. (1978) indicated that in addition to ariid catfish, the grayii species also preyed upon plotosid catfish and puffers. They also suggested that factors including innate preferences, Peduncle individual habits and availability of prey determined the selection of dietary items (prey) by sea snakes. However, experimental data on sea snake feeding habits (Voris et al. Scales of E. schistose 1978) indicate a wide range of food items in the gut probably attributed only to prey availability. However, gut contents of L. curtus revealed an unidentified clupeiform fish in a semi- digested state. Glodek & Voris (1982) and Lobo et al. (2005) indicate that L. curb's is a generalist feeder known to prey upon more than eight different fish families from different depths. Image 4. Symbiotic association between sea snake and barnacle Further, comparison of the seasonal trends of capture of (a) Eplzoite pedunculate barnacle, Octolasmis gray!! sea snakes in trawl catches (Table 3) with those of the prey (encircled) on tall of sea snake, Enhydrina schistosa organisms in the trawl catches revealed an inverse relationship (b) Enlarged view of Octolasmis gray!! with prey organisms such as catfishes (Family Ariidae), anchovies (Family Engraulidae), and clupeids (Family Clupeidae), therefore suggesting that high numbers of sea although they were found to be absent offshore, suggesting snakes had a marked effect on these groups (Fig. 2). Such shoreward migration. Shuntov (1971) indicated that sea snakes observations highlight the importance of these groups as food were widely dispersed during winter and were concentrated items of the sea snakes. Voris et al. (1978) and Lobo et al. 614 Journal of Threatened Taxa I www.threatenedtaxa.org I December 2009 I 1(12): 609.616 Sea snakes of west coast of India V.P. Padate et al.

(2005) also indicated dominance of these items in the gut before subsequent shedding of the skin. The observations content of sea snakes. On the other hand, seasonal trends in made in the present study emphasize the need for further number of sea snakes did not exhibit any particular relationship investigations of the mode of attachment and the nature of with those of other prey organisms such as portunid crabs and interactions. penaeid prawns (Fig. 2), indicating less dependence on these species as prey items. Why study sea snake ecology? In an attempt to focus on the niche level of sea snakes in the The data obtained in the present study indicate that bentho-pelagic environment, the data collected on the sea snakes approximately two snakes are incidentally trapped per trawl and the gut .contents were carefully analyzed and presented in haul. Although, this figure appears to be small, if one computes the form of a flow chart (Fig. 3) to hypothetically assess the the cumulative effect of such undesirable removal ever since trophic position of sea snakes by using a trophic scale (Pauly et the mechanization of fishing craft and trawling gear along the al. 1998) that represents the primary constituents of the marine Goan coast since 1963, it appears that such action can create food web that interact among themselves through complex prey- significant changes in the benthic coastal environment. In India, predator relationships. In this exercise it was assumed that sea snakes are protected under Section IV of the Wildlife about 90% of the energy is lost during the transfer from one (Protection) Act, 1972. However, prevention and control of trophic level to the next higher one (Odom 1959; Kozlovsky unintentional trapping of sea snakes by legislation and its 1968), with exception of certain groups like sardines and subsequent implementation is a difficult task owing to the lack anchovies that are involved in complex interactions at different of appropriate infrastructure and monitoring mechanisms along trophic levels. In such cases, the energy values were derived such an extensive coastline. Further, there is a dearth of by obtaining means of the above constituents involved in management strategies such as use of exclusion devices to avoid distinct food chains. The occurrence of sea snakes in the bentho- entry of sea snakes in the trawl net. In light of the above, the pelagic habitat coupled with their carnivorous feeding habit present study attempts to elucidate the ecological significance indicates that they act as top predators in the community, of these organisms in the coastal marine environment and feeding mainly on nektonic populations. Voris (1972) suggested recommends strict enforcement of legislation and adoption of that sea snakes use a wide range of food types and opined that appropriate management practices to avoid removal of sea the data collected on the diet of sea snakes could be used to snakes. construct a qualitative picture of the possible role of sea snakes Review of sea snake ecology from the Indian waters reveals in trophic dynamics. that available information is poor and scanty. Therefore, a holistic approach to investigate the ecology of sea snakes may Spawning be a useful tool to bring about improved understanding and Our observations on the spawning of sea snakes were enable development of effective conservation and management primarily based on dissection of individuals (N = 41) to expose plans. their reproductive systems. Among the dissected individuals one E. schitosa female, captured from the estuarine waters at 5- REFERENCES 8 In depth, contained 19 eggs (11 in the right and eight in the left uterine tube) (Image 3). The means and standard deviation Ansari, Z.A., A. Chatterji, B.S. Ingole, R.A. Sreepada, C.U. Rivonker & A.H. Parulekar (1996). Community structure and seasonal variation for diameter and weight (N = 19) of her ova were found to be of an inshore demersal fish community at Goa, West Coast of India.

4.20±0.05 cm and 14.09±1.00 g, respectively. Voris and Jayne Estuarine, Coastal and Shelf Science 41: 593 -610. (1979) stated that sea snakes give birth to offspring during the Ansari, Z.A., R.A. Sreepada, S.G. Dalai, B.S. Ingole & A. Chatterji post-monsoon season in the inshore waters of Malaysia and (2003). Environmental influences on the trawl catches in a bay further suggested that clutch size increases with the size of the estuarine system of Goa. Estuarine, Coastal and Shelf Science 56: 503- 515. female and can reach thirty or more. However, Karthikeyan et Burns, G., & H. Heatwole (1998). Home range and habitat use of the al. (2008) reported a maximum of three to five clutches in Olive Sea Snake, ilipysurus laevis, on the Great Barrier Reef, Australia.

Hydrophis cyanocintus females and opined that clutch size Journal of Herpetology 32: 350 - 358. increases with the size of the females in this species. Dunson, W.A. (1975). Adaptation of sea snakes, pp. 3-19. In: Dunson, W.A. (ed.). The Biology of Sea Snakes. University Park Press, Baltimore, Symbiotic associations xi+530pp. Fernando, S.A. (2008). Monograph on Indian barnacles - Marine Benthos Sea snakes (N = 5) were associated with barnacles (Image 02. Ocean Science and Technology Cell, Cochin University of Science 4a), a constituent of inter-tidal rocky zones and which have a and Technology, Kochi, 199pp. cementing gland for attachment. In the present study, these Fischer, W., & P.J.P. Whitehead (1974). MO Species Identification Sheets individuals were found to be attached to the surface of the sea for Fishery Purposes. Eastern Indian Ocean (fishing area 57) and Western snakes. These barnacles were identified as Central Pacific (fishing area 71) Volume I-IV. Food and Agricultural Octolasmis grayii Organization, Rome. (Image 4b). This is the first report of such an association among Fry, G.C., D.A. Milton & T.J. Waasenberg (2001). The reproductive sea snakes and barnacles from the tropical inshore waters of biology and diet of seasnake by-catch of prawn trawling in northern Goa. Although, Jeffries & Voris (1996) remarked that different Australia: characteristics important for assessing the impact on types of symbiotic associations prevail among marine population. Pacific Conservation Biology 7: 55 -73. organisms, there are no specific studies that elucidate the nature Glodek, G.S. & H.K. Voris (1982). Marine snake diets: prey composition, diversity and overlap. Copeia 1982: 661 - 666. of this association. The present observations suggest that it Goa, Daman and Diu Marine Fishing Regulation Rules (1982). could be parasitic, as the stalked barnacle (Octolasmis grayii), Notification 2-2 - 81 - FSH. Government of Goa, Daman and Diu, attach to these reptiles following shedding of the skin, Forest and Agriculture Dept, Panaji, Goa, India. primarily on large-sized individuals and they get detached Heatwole, 11. (1997). Marine snakes: are they a sustainable resource?

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Wildlife Society Bulletin 25: 766 - 772. Acknowledgements: The authors take this Ireatwole, II. (1999). Sea Snakes. University of New South Wales Press Limited, Sydney, 149pp. opportunity to express their gratitude to the Ballast Hyslop, E.J. (1080). Stomach contents analysis - a review of methods and their application. Water Management Programme, India executed by National Institute of Oceanography, Dona Paula, Journal of Fish Biology 17: 411 - 129. Goa for Directorate General of Shipping, Ministry of Jeffries, W.B. & ILK. Voris (1996). A subject-indexed bibliography of the symbiotic barnacles Shipping, Government of India. Further, we express of the genus Octolastnis grayii, 1825 (Crustacea: Cirripedia: Poecilastnatidae). The Raffles Bulletin our sincere gratitude to Dr. H. Heatwole and the of Zoology 44: 575 - 592. anonymous referee for their constructive criticism Karthikcyan, R., S. Vijayalaksmi & T. Balasuhramanian (2008). Feeding and parturition of that enabled Improvement in the quality of the female annulated sea snake Ilydrophis cyanocincess in captivity. Current Science 94: 660-664. manuscript. We would also like to thank the trawl Kozlovsky, D.G. (1068). A critical evaluation of the trophic level concept. I. Ecological efficiencies. owners and the crew of *Jesus Bless" for permitting us to carry out on-board sample collection. Ecology 49: 4S - 60. Kulkarni, S.S., C.U. Rivenker & U.M.X. Sangodk: r (2003). Role of meiobenthic assemblages in detritus based food chain from estuarine environment of Goa. Indian Journal of Fisheries 50:

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marine snakes along the coast of Goa, Western India. Hamadryad 29: 89 -93. Lobo, A.S., K. Vasudevan & B. Pandav (2005). Trophic ecology of Lapentis curtus (Hydrophiinae) along the Western Coast of India. Copeia 2005: 636-640. Odum, E.P. (1959). Fundamentals of Ecology. 2nd edition. Saunders, Philadelphia, 546pp. Pauly, D., V. Christensen, J. Dalsgaard, R. Freese & J.F. Torres (1098). Fishing down marine

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Rasmussen, A.R. (2001). Sea Snakes, pp. 3987 -1003. In: Carpenter, K.E. & V.H. Niem (eds.). FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 6. Food and Agricultural Organization, Rome. Reynolds, R.1'. & C.V. Pickwell (1984). Records of the yellow-bellied sea snake, Pelamis platnrus,

from the Galapagos Islands. Copeia 1984: 786 - 789.

Shuntov, V.P. (1971). Sea snakes of the north Australian shelf (in Russian). Ekologiya 2: 65 - 72. Tu, A.T. (1974). Sea snake investigation in the Gulf of Thailand. Journal of Herpetology 8: 201- 210. Venkataraman, K. & M. Wafar (2005). Coastal and Marine biodiversity of India. Indian Journal of

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ocean communities. Journal of the Marine Biological Association of India 14: 429 - 442. Voris, IT.K. (1985). Population size estimates for a marine snake (Enhydrina schistose) in Malaysia.

Copeia 1085: 955 - 961. Voris, H.K. & B.C. Jayne (1979). Growth, reproduction and population structure of a marine

snake, Enhydrina schistose (Hydrophiidae). Copeia 1979: 307 - 318. Voris, ILK. & MIL Voris (1983). Feeding strategies in marine snakes: an analysis of evolutionary,

morphological, behavioral and ecological relationships. American Zoologist 23: 411 -425. Voris, ILK., 11.H. Voris & C.B. Liat (1978). The food and feeding behavior of a marine snake,

Enhydrina schistose (Ilydrophiidae). Copeia 1978: 131 - 146. Wass ,..hberg, T.J., J.P. Salini, H. Heatwole & J.D. Kerr (1994). Incidental capture of sea snakes (11ydrophiidae) by prawn trawlers in the Gulf of Carpentaria, Australia. Australian Journal of

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skin of sea snakes, pp. 251 -265. In: Dunson, W.A. (ed.). The Biology of Sea Snakes. University Park Press, Baltimore, xi+SSOpp.

616 Journal of Threatened Taxa I www.threatenedtaxa.org I December 2009 I 1(12): 609-616 Marine Biodiversity Records, page s. of 6. ,t Marine Biological Association of the United Kingdom, 2010 dOl:10.101261755267210000151; Vol. 3; en; 2010 Published online A note on the occurrence of reef inhabiting, red-bellied yellow tail fusilier, Caesio cuning from outside its known geographical array

VINAY P. PADATE ' , CHANDRASHEKHER U. RIVONKER ' AND A. CHANDRASHEKAR ANIL2 'Department of Marine Sciences, Goa University, Goa 403 206, India, .'National Institute of Oceanography (CSIR), Dona Paula, Goa 403 004, India

The reef inhabiting fusilier, Caesio coning, characterized by pinkish abdomen and yellow caudal fin exhibits shoaling behaviour and is known to be geographically distributed from the Gulf of Mannar to the Vanuatu Islands in the South Pacific. The present observation is evidence for the first record of this species off Goa, west coast of India, to be outside its known geographical array. The bay-estuarine and coastal region of Goa influenced by south-west monsoon receives nutrient-rich water during up welling. Hence the region is highly productive and supports diverse marine assemblages. Moreover, the present observation reveals very low numbers (N = 4) of this species in the light of the trawl hauls (N = 92) with a core sampling effort of 146 hours, wherein it was found to occur only in two hauls. The paper also provides detailed insight into the description of C. cuning from this region with orthodox taxononpc methods including morphometry and meristic counts.

Keywords: reef inhabiting, Caesio cuning, first record, geographical array, Goa, west coast of India

Submitted 30 January 2009; accepted 20 November 2009

INTRODUCTION geographical locale in the coastal waters of Goa along the west coast of India. The zoogeographical range of a species refers to the distri- bution of a species, either horizontal or vertical, largely determined by the pervasiveness of certain environmental MATERIALS AND METHODS anomalies and biotic components that favour establishment of the species, and whose boundaries are defined by gradual The present study area encompasses the Mandovi-Zuari estu- dilution or abrupt termination of these components. Coastal arine complex (Figure 1) and the adjacent fishing grounds up to ecosystems being dynamic are rich in diverse assemblages 20 m depth that are subjected to increased levels of exploitation with estimates of about 80% of marine fauna inhabiting by commercial trawl operations. The potential fishing grounds these areas (Ray, 1988, 1991; Anon, 1989) with highly overlap- located between 15"29'N and 15"34'N latitudes and between ping boundaries that are influenced by complex relationships 73"40"E and 73"51'E longitudes were selected for intensive among varying niche's in the marine environment. These sampling. During the present study period (May, 2005 to coastal areas also support rare fish species (Engel & Kvitek, May, 2008), intensive sampling comprising 92 trawling oper- 1998; Froese & Pauly, 2008) predominantly belonging to ations with a total effort of 146 hours was undertaken in the two finfish orders, namely Perciformes (perch-like fish) and estuarine and offshore areas of Goa, central west coast of Scorpaeniformes (scorpion fish, rock fish and sting fish) India to assess the diversity and total community structure of (Froese & Pauly, 2008). Among the perch-like fish, the fusi- demersal fish fauna. Trawl-nets with mesh sizes of 15 mm liers belonging to the family Caesionidae are particularly (mouth-end) and 9 mm (cod-end) were towed approximately unique with regards to evolutionary trends that display tran- at a speed of 2 knots (4 km. h - '). The catch obtained was sition from benthic carnivorous to semi-pelagic planktivorous sorted into five sub-samples. Uncommon (or rare) specimens behaviour (Carpenter, 1988). were picked out, put on ice and sent to the laboratory for The inshore shelf waters of Goa, central west coast of India detailed examination. In addition, samples were obtained by up to 20 in depth, are subjected to anthropogenic inputs such operating beach seine along Betim (15'30'18"N 73°49'52"E) as commercial trawling (Ansari et al., 1995, 2003) and mari- during December, 2005 and twice in the vicinity of time activities (Anil et al., 2002) that pose a potential threat Mormugao Port Trust (15'24'36"N 73"48'56"E) during to coastal ecosystems and the resident biota. The present December, 2005 and September, 2006 to assess the occurrence paper describes the new record of occurrence of rare fusilier of demersal species in the estuarine embayment as these were species, Caesio cuning (Bloch, 1.791) found outside its inaccessible to demersal trawlers. At the laboratory, a detailed morphological study of the

Corresponding author: biological specimens was carried out using identification C.U. Rivonker keys (Fischer & Whitehead, 1974; Talwar & Kacker, 1984; [email protected] Froese & Pauly, 2008). ▪

VINAV 0. PADATE ET Al..

Area Expanded

vow MV Hatt Pnrrcax

•gf.or if svirws ,, ,I,e,-;1"r ■

Fig. r. Map of the study area indicating sampling sites and location of collection of Caesio coning.

RESULTS Family CAESIONIDA14

Ascending maxillary process is completely separated from MATERIAL EXAMINED the pre-maxilla (Carpenter, 1988). In other related perciform Two unsexed specimens; RP 1, 6.08 cm SI. and RP-2, 5.21 cm fish, particularly snappers (family Lutjanidae), the ascending SL (co-ordinates: 15 '30'43.1"N 73 -45'40.1"E to 15“32'28.1"N maxillary process is confluent with the pre-maxilla 73"45'9.4"E; water depth: i i - 14 m). Coll. V.P. Padate, (Carpenter, 1988). The above morphological character is an Ma rch 2007. adaptation towards a planktivorous feeding habit. Two unsexed specimens; RP-3, 6.89 cm SL and RP-4, The taxonomic identification of these specimens up to the 6.11 cm SL (co-ordinates: 152943.2"N 73' 44'39.7"E to species level was carried out using identification keys provided

1 5 33'15.2"N 73 42'41.6"E; water depth: 6 -7 m). Coll. V.P. by Carpenter (1988) and Froese & Pauly (2008). Padate, 25 March 2007. Details of morphometric measurements of fish specimens Genus Caesio Lacepede, 18oi along with their mean and standard deviation are provided A single post-maxillary process; posterior end of maxilla in Table I. Reference voucher sample is deposited at the blunt, its greatest depth posterior to end of pre-maxilla Marine Biology Laboratory, Department of Marine Sciences, (Carpenter, 1988). Goa University. Caesio tuning (Bloch, 1791) (Figures 2 - 7)

Table 1. Details of morphometric measurements (in cm) of Caesio coning specimens (N =. 4) collected along the coast of Goa. Dorsal region of body from tip of snout to anterior spinous • portion of dorsal fin greyish yellow, mid-lateral portion of Serial Morphological Specimen Mean ± SD body pale-white (Figure 2); caudal fin yellow, lacking No. parameter FP 1 RP-2 RP•3 RP-4

Total length 8.48 7.49 10.05 8.56 8.65 + 1.06 2. Fork length 6.99 6.57 8.38 7.42 7.34 ± 0.78 1. Standard length 6.08 5.11 6.89 6.11 6.07 4- 0.69 Body depth 2.34 1 .91 IN 2.27 2.29 t 0.30 Pre-dorsal length 2.28 2.07 2.53 2.32 2.30 + 0.19 6 Pre-anal length 4.18 NA 4.84 4.33 4.45 ± 0.35 7. Head length z.oz NA .2.30 2.1,3 2.17 t 0.17 8. Snout length 0.39 0.42 0.44 0.38 0.41 ± 0.03 y. Eye diameter 0.74 0.01 o.88 0.87 0.78 ± 0.13

RP, site of sample collection in the vicinity of the MV 'River Princess' Fig. a. Distinguishing characters and colour of a fresh specimen of Caesio grounded off the coast of Goa. CI. I A NOTE ON THE OCCURRENCE OF REEF INHABITING 3

Fig. 3. Caudal fin of Caesio dining indicating characteristic yellow coloration. prominent blackish markings (Figure 3); ventral region of body from cheeks to lower portion of caudal peduncle, pec- toral, pelvic and anal fins light pink (Figure 4). Supra-temporal band of scales of either side confluent at dorsal midline (Figure 5); dorsal fin with ten spines and i5 Fig. 5. Supra-temporal band of scales in Caesio caning. (A) Present specimen; soft rays, its spinous portion covered with scales up to half (13) diagrammatic representation of supra-temporal scales of C. caning its greatest height (Figure 6); anal fin with three spines and provided by Carpenter (1988).

II soft rays, covered with scales (Figure 7); about 4 - 5 scale rows on cheek; 48 -51 scales in lateral line. The data on morphometry of present specimens was compared with existing literature. FishBase, the world-wide geo- graphical information system on finned fish (Froese & Pauly, 2008) reveals that Caesio cuning grows to a size of 6o cm total length (TL). However, the present specimens (N = 4) ranged between 7.49 cm and 10.05 cm TL, thus confirming the capture of juvenile/young individuals of this species.

DISCUSSION

The existing data on the geographical distribution of Caesio cuning (Figure 8) suggest that this species inhabits the Indo-Pacific region from the Gulf of Mannar to the Vanuatu Fig. 6. Dorsal fin of Caesio curling indicating scaled portion and characteristic fin count. Islands in the South Pacific (Carpenter, 1988; Froese & Pauly, 2008). Carpenter (1988) attributed the geographical distri- bution of this species to availability of appropriate habitat Subsequently, the efforts of the three-year study led to the and adequate ecological parameters that favour the occurrence reporting of C. curling for the first time off Goa and along of this species. Published literature on the commercial finned the entire west coast of India. It has been well documented fish of the Indian coast (Talwar & Kacker, 1984) indicates earlier (Carpenter, 1988) that this species and its congeners that C. cuning occurs along the Andaman coast which falls occur in coastal reef areas. The present specimens were within the known zoogeographical range of this species. obtained from bottom trawl catches in the vicinity of the However, FishBase (Froese & Pauly, 2008) indicates the possible expansion of occurrence of this species to the adjacent waters. Published literature along the Goa coast (Prabhu & Dhawan, 1974; Ansari et al., 1995, 2003) provides information on the total demersal fish corhmunity structure with major emphasis on commercial finfish species. However, this species is not men- tioned in available literature from the present study area. In view of the above, our observations primarily focus on the occurrence of rare species along the coast of Goa.

3 spines

Fig. 4. Ventral portion of Caesar caning indicating characteristic pink coloration. Fig. 7. Anal fin of Caesio caning indicating characteristic fin count. 4 VINAY P. PADATE ET AL.

Fig. 8. M. p of the world indicating geographical distribution of Caesio cuning a. Known zoogeographical range of Caesio cutting; a, Possibility of occurrence of Caesio cunin,v; GOA, site of present collection.

grounded ore-carrier, MV 'River Princess' off the Candohm - adjacent Mandovi estuary and Aguada Bay (Ansari et al., Sinquerim shore. The MV 'River Princess' grounded off the 1995). Kessarkar et al. (2009) reported two seasonal peaks coast of Goa in 2000 (Ingole et al., 2006) and might have in suspended particulate matter (SPM) concentration created an artificial reef-like habitat, thereby attracting (zo mg. 1 - ' and 19 mg. 1 - during June-September and several reef inhabitants from adjacent reefs (Qasim & February-April, respectively) from the Mandovi estuary Wafar, 1979) and further facilitated recruitment of their and attributed these to interactions between strong seasonal larval and juvenile forms (Arena et al., 2007). FishBase winds, wind-induced waves and tidal currents in the (Froese & Pauly, 2008) reveals that C. cuning grows to a size estuary. It is probable that wind-induced waves, tidal currents, of 60 cm TL. However, the present specimens (N = 4) along with the resultant turbidity in the water column, might ranged between 7.49 cm and 10.05 cm TL, thus confirming have induced the migration of C. cuning juveniles towards the capture of young stages. Secondly, existing literature indicates bay-estuarine waters. Carpenter (1988) suggested that C. the presence of partially submerged rocky reefs off the Aguada cuning prefers turbid waters, thus corroborating the present Hill, that are inaccessible for trawl-net operations (Qasim & observations. Further, Leis & Carson-Ewart (2003) suggeu Wafar, 1979), thus suggesting suitability of habitat in this the orientation of pelagic larvae of certain reef-fish iii response area. Moreover, the above region has nutrient-rich turbid to solar or reef-based cues. The present observations reveal waters owing to transport of suspended material from the that the prevalence of similar ecological conditions might have enabled the younger population of C. cuning to

35 swine —4— migrate to a coastal -estuarine embayment. 10tri Caesio cuning inhabits the eastern Indian Ocean and western Pacific Ocean regions (Carpenter, 1988) which are 30 influenced by hydrographic parameters (temperature, salinity

td and dissolved oxygen) characteristic of tropical oceans 25 (Moore, 1972). Hydrographic parameters measured during the present study were observed to be similar and indicated 36 clear seasonal variations at to in depth (Figure 9), however, the value was found to be comparatively higher at the time 34 of collection of C. cuning. The study area experiences a seaso- 5, 32 nal upwelling phenomenon (Ansari et al., 1995) rendering the region highly fertile in terms of primary productivity (13.12 30 to 14.21 mg c.nt '.11 -1 , Krishna Kumari et al., 2002). Such 200 high productivity sustains high zooplankton biomass

9- • (Qasim & Sen Gupta, 1981). As C. culling is a zooplankton 1 160 feeder (Carpenter, 1988), it is pOssible that sehoolS of this species might have migrated to the above region 100 . to forage upon the abundant zooplankton. 1 so Furthermore, the intensive sampling effort covering a Jan Feb Mn, Apr May Jun Jul Aug Sep Oct Nov Dec wider coastal region through use of modern fishing vessels Month and gear that provide access to submerged rocky patches in

Mg. 9. Monthly variations in hydrographic parameters at surface and to in coastal inshore embayment may have enabled entrapment of depths during 2007. this species. However, it must be noted that in the present A NOTE ON THE OCCURRENCE OF REEF INHABITING 5 study, only four specimens were obtained in the light of the Anon (1989) Loss of biological diversity: a global crisis requiring inter- intensive sampling (N = 92) with a core fishing effort of 146 national solutions. Washington, DC: National Science Foundation. hours over a period of three years excluding monsoonal Ansari Z.A., Chatterji A., Ingole B.S., Sreepada R.A., Rivonkar C.U. periods (June-August), wherein it was found to occur only and Parulekar A.H. (1995) Community structure and seasonal in two hauls. The present observations contradict earlier variation of an inshore demersal fish community at Goa, west coast observations (Carpenter, 1988) that suggest a shoaling behav- of India. Estuarine, Coastal and Shelf Science 41, 593-61o. iour, indicating either these individuals might have strayed Ansari Z.A., Sreepada R.A., Dalai S.G., Ingole B.S. and Chatterji A. away from the larger shoal and got trapped in the trawl-net (2003) Environmental influences on the trawl catches in a bay estuar- over the soft substratum or might have not been able to estab- ine system of Goa. Estuarine, Coastal and Shelf Science 56, 503-515. lish as a large shoal largely due to habitat complexity. Moreover, the study area is a potential fishing ground and Arena P.T., Jordan L.K.B. and Spieler R.E. (2007) Fish assemblages has been intensively exploited by demersal trawlers peren- on sunken vessels and natural reefs in southeast Florida, USA. Hydrohiologia 58o, 157-171. nially (with the exception of a 75-day monsoon ban) ever since the initiation of mechanized fishing in 1963 and its Carmen M.A. and Ablan A. (2006) Primer note: microsatellite loci further expansion during the subsequent decades. Such long- for studies on population differentiation and connectivity of the term intensive trawling activity may lead to changes in the red-bellied yellow tail fusilier, Caesio cuning (Caesionidae). structure and composition of the resident demersal commu- Molecular Ecology Notes 6, 170-572.' nities and benefit other species groups as reported earlier Carpenter K.E. (1988) FAO Species Catalogue. Voltme 8. Fusilier fishes of from the Gulf of Thailand (Longhurst & Pauly, 1987). the world—an annotated and illustrated catalogue of caesionid species known to date. Rome: FAO.

CONCLUSION Engel J. and Kvitek R. (1998) Effects of otter trawling on a benthic com- munity in Monterey Bay National Marine Sanctuary. Conservation Biology 12, 1204-1254. Our observations suggest that this is the first new record of Caesio cuning off Goa along the entire west coast of India that Froese R. and Pauly D. (eds) (2008) FishBase world wide web electronic lies outside its known geographical array. The present study, publication. http://www.fishbase.org version (07/2008). through intensive sampling effort of 146 hours over three Fischer W. and Whitehead P.J.P. ( 1 974) FAO Species Identification Sheets years, attempted to highlight the occurrence of the rare for fishery purposes, eastern Indian Ocean (fishing area 57) and western species, C. curling. The occurrence of this species in low central Pacific (fishing area 71). Volume 1. Rome: FAO. numbers could primarily be due to sampling gear and local bathymetry in the vicinity of the grounded ship that probably Ingole B.S., Sivadas S., Goltekar R., Clemente S., Nanajkar M., Sawant (2.006) Ecotoxicological effect reduced the chances of entrapment of this species in the trawl- R., D'Silva C., Sarkar A. and Ansari Z. of grounded MV River Princess on the intertidal benthic organisms off net. Furthermore, it appears that the selection of gear with Goa. Environment International 32, 284-291. increased efficiency in such reef dominated habitats might yield more individuals from such areas. It is suggested that Kessarkar P.M., Purnachandra Rao V., Shynu R., Ahmad I.M., Mehra further detailed ecological studies on this species might P., Michael G.S. and Sundar D. (2009) Wind-driven estuarine turbid- provide more detailed information and explain its occurrence ity maxima in Mandovi Estuary, central west coast of India. Journal and various biological Vrocesses in such habitats. Moreover, mol- of Earth System Science 118, 369-377. ecular studies of this species using genomic DNA would further Krishna Kumari L., Bhattathiri P.M.A., Matondkar S.G.R. and Julie enhance the understanding of its population stock structure and John (2002) Primary productivity in Mandovi-Zuari estuaries in connectivity with existing populations (Carmen & Ablan, 2006). Goa. Journal of the Marine Biological Association of India 44, 1-13. Leis J.M. and Carson-Ewart B.M. (2003) Orientation of pelagic larvae of coral-reef fishes in the ocean. Marine Ecology Progress Series 252, ACKNOWLEDGEMENTS 239-253. The authors take this opportunity to express their gratitude to Longhurst A.R. and Pauly D. (eds) (1987) Ecology of tropical oceans. San the Bollast water Management Programme, India executed by Diego, CA: Academic Press Inc. National Institute of Oceanography, Dona Paula, Goa for Moore H.B. (1972) Aspects of stress in the tropical marine environment. Directorate General of Shipping, Ministry of Shipping, In Russell F.S. and Yonge M. (eds) Advances in marine biology. Government of India. We also thank the owner and crew of Volume io. London: Academic Press Inc., pp. 217-269. the fishing trawler 'Jesus Bless' for assisting in the intensive sampling programme. We would also like to take this opportu- Prabhu M.S. and Dhawan R.M. (1974) Marine fisheries resources in the 20 and 40 metre regions off the Goa coast. Indian Journal of Fisheries nity to express thanks to Mr Venkat Krishnamurthy for the specimen photography and artwork included in this manuscript. 47, 40- 53. This is a joint contribution of Goa University and National Qasim S.Z. and Sen Gupta R. (1981) Environmental characteristics of the Institute of Oceanography (NIO contribution no. 4661). Mandovi-Zuari estuarine system in Goa. Estuarine, Coastal and Shelf Science 13, 557-578. Qasim S.Z. and Wafar M.V.M. (1979). Occurrences of living coral at REFERENCES several places along the west coast of, India. Mahasagar NIO, Goa t 2, 53-58. Anil A.C., Venkat K., Sawant S.S., Dileepkumar M., Dhargalkar V.K., Ramaiah N., Harkantra S.N. and Ansari Z.A. (2002) Marine Ray G.C. (1988) Ecological diversity in coastal zones and oceans. bio-invasion: concern for ecology and shipping. Current Science 83, In Wilson E.O. and Peter F.M. (eds) Biodiversity. Washington, 214-218. DC: National Academy Press, pp. 36-5o. 6 VINAY P. PADATE. ET AL.

Ray G.C. (1991) Coastal zone biodiversity patterns. Bioscience 41, Correspondence should be addressed to: 490- 49 8. C.U. Rivonker Department of Marine Sciences and Goa University Talwar P.K. and Kacker R.K. (1984) Commercial sea fishes of India. Goa 403 206, India Calcutta: Zoological Survey of India. email: [email protected] Marine Biology Research, 2010; 6: 579-590

ORIGINAL ARTICLE

A new species of portunid crab of the genus Charybdis (De Haan, 1833) (Crustacea: Decapoda: Brachyura) from Goa, India

VINAY P. PADATE', CHANDRASHEKHER U. RIVONKER1 *, A. CHANDRASHEKAR ANIL2, SUBHASH S. SAWANT2 & VENKAT KRISHNAMURTHY2

'Department of Marine Sciences, Goa University, Goa, India; 2National Institute of Oceanography (CSIR), Dona Paula, Goa, India

Abstract A new species of portunid crab, Charybdis (Charybdis) goaensis sp. nov. is described from Goa, west coast of India. It differs from its closest congener, Charybdis (Charybdis) philippinensis in possessing bluntly triangular median frontal teeth, wide U-shaped notches separating the sub-median and lateral frontal teeth, wide notches separating antero-lateral teeth, and granular tuberosity on cardiac region of carapace. A taxonomic key to 43 species of the sub-genus Charybdis including the newly described species is also provided.

Key words: Charybdis (Charybdis) goaensis, Crustacea, Decapoda, India, new species, Portunidae

Introduction with detailed descriptions of 33 species and 7 varieties. Subsequent taxonomic work on portunid Swimming crabs (Family Portunidae Rafinesque- crabs by Stephenson (1972) provided the most Schmaltz, 1815) are crustaceans which inhabit extensive taxonomic key to the genus Charybdis. sub-tidal, estuarine and offshore waters, and are Studies pertaining to systematics of crabs of the widely distributed across the Indo-West Pacific region genus Charybdis from the Indian Ocean region were (Stephenson 1972). Charybdis De Haan, 1833, is the initiated by the 'Discovery' expedition (Stephenson second largest genus within the sub-family Thalami- & Rees 1967a) which revealed one previously known tinae Paulson, 1875, with 60 species (Thalamita species, C. smithii. This was followed by six expedi- Latreille, 1829 comprises 89 species) (Ng et al. tions in the Western Indian Ocean region (`RN 2008) distributed among four sub-genera (Charybdis Akademic Petrovsky', `R/V Odissey', `R/V Akademic De Haan, 1833; Goniohellenus Alcock, 1899; Gonio- Kurchatov', `R/V Meteor', `R/V Professor Mesyat- neptunus Ortmann, 1893; Goniosupradens Leene, sev', `R/V Vitiaz') that found nine species in 1938). which two (C. crosnieri and C. meteor) were new to For the past two centuries, naturalists have con- science (Neumann & Spiridonov 1999; Spiridonov tributed immensely to the taxonomy of the genus & Turkay 2001). Subsequently, the `Anambas' Charybdis (De Haan, 1833) through extensive study expedition in the Eastern Indian Ocean region of brachyuran fauna collected from deep-sea and revealed one previously known species, C. truncates coastal expeditions as well as from museum collec- (Yeo et al. 2004). Recently, Apel & Spiridonov tions. Leene (1938) carried out the first major (1998) dealt with portunid crabs of the Arabian taxonomic work on crabs of the genus Charybdis. Gulf, the Gulf of Oman, southern Oman and She introduced two new sub-genera, namely Gonio- Pakistan. supradens and Gonioinfradens, to the existing three A review of published literature reveals that out of (Charybdis, Goniohellenus and Gonioneptunus) along 43 extant species of the sub-genus Charybdis, 35

*Correspondence: Chandrashekher U. Rivonker, Department of Marine Sciences, Goa University, Goa 403 206, India. E-mail: curivonker0t,?gmail.com Published in collaboration with the University of Bergen and the Institute of Marine Research, Norway, and the Marine Biological Laboratory, University of Copenhagen, Denmark

(Accepted 21 October 2009; Published online 4 August 2010; Printed 14 October 2010) ISSN 1745-1000 print/ISSN 1745-1019 online © 2010 Taylor & Francis DOI: 10.1080/17451000903505608 580 V. P Padate et al. species inhabit estuarine and offshore shallow waters Terminology used in the morphological descrip- (Alcock 1899; Leene 1938; Ward 1941; Wee & Ng tion of the new species follows Wee & Ng (1995). In 1995), 3 inhabit deep-sea as well as shallow coastal addition, terminology describing thoracic sternum environments (Stephenson & Rees 1968; Spiridonov and abdominal cavity follows Ng (2000). & Ti1rkay 2001) and 5 species inhabit deep-sea Thirty-nine trawling stations were selected in the environments (Moosa 1995; Spiridonov & Tiirkay estuarine and offshore areas of Goa, west coast of 2001). India (Figure 1) to assess the diversity and total In addition to the extant species of the genus community structure of the demersal fauna. Among Charybdis, recent published reports (Hu & Tao these, sampling was carried out at six stations from 1979, 1985, 1996, 2000; Morris & Collins 1991) the estuaries during May and December 2005, and described five extinct species (C. kilnzeri Hu, 1984; September and October 2006, along the navigational C. monsoonis Hu & Tao, 1985; C. feriatus anzenia channel off Mormugao Port Trust (15°25'N, Hu & Tao, 2000; C. gigantica Hu & Tao, 2000; C. 73°47'E) region and the southern region of Mormu- feriata bruneiensis Morris & Collins, 1991) from East gao Bay (between 15°24'N, 73°48'E and 15°27'N, and South-east Asia based on diagrammatic recon- 73°51'E). The offshore samples were obtained struction of fossilized remnants of crabs. However, from January to April 2006 and December 2006 to these species were not compared with the extant March 2007 between 15°29'6'N and 15°33'15.2'N species in view of the ambiguity in their description. and between 73°40'6'E and 73°51'1 l'E. Trawl nets A new species of swimming crab, Charybdis with mesh sizes of 15 mm (mouth end) and 9 mm (Charybdis) goaensis sp. nov. from the estuarine and (cod end) were towed approximately at a speed of offshore waters of Goa, west coast of India is 2 knot (4 km h — '). The sampling duration was described here (Figure 1). Further, the new species 1-3 h in the offshore region, whereas in the estuaries is compared with all the existing congeners and an it lasted for a maximum of 1 h. The catch obtained updated identification key incorporating 43 species was sorted into five sub-samples. Uncommon (or of the sub-genus Charybdis is provided. rare) specimens were picked out, put on ice and sent to the laboratory for detailed examination. In addi- tion, samples were obtained by operating beach Material and methods seine (1 h operation) along Betim (15°30'18'N, Abbreviations 73°49'52'E) during December 2005 and two in the vicinity of the Mormugao Port Trust (15 °24'16'N, The following abbreviations are used: CL, carapace 73°48'56'E) during December 2005 and September length; CW, extreme width of carapace; LC, length 2006 (Figure 1). of left cheliped; RC, length of right cheliped; At the laboratory, morphological characteristics of OD, orbital diameter; IS, inter-orbital space/frontal the crabs were recorded by camera lucida diagrams width; GI, male gonopod/first pleopod. and photographs. Seven morphometric parameters

Urea kmanded

Figure 1. Map illustrating the site of collection of Charybdis (Charybdis) goaensis sp. nov. specimens. New species of portunid crab 581 were measured using vernier callipers with an Charybdis (Charybdis) goaensis sp. nov. accuracy of 0.01 mm. The three ratios (CW/CL, OD/IS and IS/CW) were derived. Material examined In the case of chelipeds, the measurement in- Holotype: male (designated here GUMS-1), 29 cluded segments namely, dactylus, propodus, carpus September 2006, CL 12.80 mm, CW 24.32 mm, and merus. Subsequently, gonopods were removed Zuari Estuary, Goa, central west coast of India from the paratype male for scanning electron micro- (between 15°24'41.2'N, 73°48'56.6'E and 15°24' scope (SEM) photography to ascertain the identity 41.9'N, 73°51'11.0'E), 6-7 m depth, demersal and distinctiveness of the species. In addition, trawl. camera lucida diagrams of the ventral surface of Paratype: Zuari Estuary, Goa: one mature female male and female specimens were drawn to illustrate (GUMS-2), CL 15.10 mm, CW 24.52 mm; one the thoracic sternites and the abdominal cavity of male (GUMS-3), CL 21.88 mm, CW 34.08 mm; male and female specimens, respectively. The speci- one male (GUMS-4), CL 14.02 min, CW mens were stored in 5% buffered formalin (buffered 25.10 mm; one immature female (GUMS-5) CL with hexamethylene tetramine to prevent fragment- 11.92 mm, CW 22.12 mm (location and date of ing of appendages) solution in transparent plastic collection same as holotype); offshore fishing bottles. These are deposited at the Marine Biology grounds, Goa: one ovigerous female (GUMS-6) laboratory, Department of Marine Sciences, Goa CL 19.40 mm, CW 29.92 mm, offshore trawl catch, University. Goa, India (between 15°29'43.2'N, 73°44'39.7'E and 15°33'15.2'N, 73°42'41.6'E), 11 March 2007. Taxonomy

Family Portunidae Rafinesque-Schmaltz, 1815 Diagnosis Genus Charybdis De Haan, 1833 Carapace evenly tomentose/pilose, with the excep- Charybdis, De Haan 1833, pp. 9-10 (type species: tion of transverse ridges and swellings. Median Cancer sexdentatus Herbst, 1783 (a junior subjective frontal teeth bluntly triangular, sub-medians and synonym of Cancer feriatus Linnaeus, 1758)). lateral frontal teeth separated by wide U-shaped notch. Orbits with strong dorsal inclination; major The genus Charybdis is characterized by the orbital diameter two-fifths to nearly half the length of fronto-orbital border being decidedly less than the inter-orbital space. Basal antennal joint with the greatest width, of the carapace; antero-lateral smooth granular ridge. Antero-lateral teeth six in borders oblique and arched, cut into six or seven number, first slightly notched, sixth greater than teeth (Sakai 1976; Wee & Ng 1995; Figure 2A); twice the length of the preceding five and laterally G1 tubular, curved or bent in distal part, tip directed; teeth separated by wide notches. Transverse usually elongated or truncated, with spines on ridges of carapace distinctly granular, two proto- lateral surfaces and marginal spinules inside open- gastric, one meso-gastric, a medially divided meta- ing, along ventral and dorsal margins (Apel & gastric, and epibranchials along the level of the last Spiridonov 1998; Figure 2B). antero-lateral teeth; no transverse ridges beyond the

A Fronto-orbital margin B

Distally bent G1 Spines on \-1 -- Six antero-lateral lateral margin teeth

f

•--

Carapace width

Figure 2. Generalized schematic diagram of Charybdis indicating distinguishing morphological characters. A, Dorsal surface of carapace (camera lucida diagram). 13, GI . 582 V P Padate et al. level of the last antero-lateral teeth; prominent Posterior margin of carapace slightly curved, forms a tuberosity on cardiac region covered with densely curve with the postero-lateral margins of carapace. spaced small granules. Chelipeds subequal in both Transverse ridges distinctly granular, two pro- sexes, rough, unevenly tomentose; merus with two to-gastric, one meso-gastric, a medially divided well-developed spines and five to six granules on meta-gastric, and epibranchials along the level of anterior margin; manus tumid, five-costate, with the last antero-lateral teeth; frontal ridge obsolete. five spines on upper surface, two on distal margin No transverse ridges beyond the level of the last immediately posterior to articulation with dactylus antero-lateral teeth; prominent tuberosity on cardiac possess blunt tips; dactylus smooth, strong and region covered with densely spaced small granules slender with curved tips, longer than manus. Propo- (Figure 3A,B). Surface of transverse ridges and dus of natatory leg smooth, merus bears prominent tuberosities more or less devoid of tomentum. spine on postero-distal margin. Sixth abdominal Chelipeds sub-equal in both sexes (LC 29.30 + segment in male broader than long, its sides parallel 7.36 mm; RC 29.04 ± 8.51 mm), rough, tomentose, over less than half their lengths and gradually a little less than twice the carapace length (Figure 3E). converge distally. GI L-shaped, its distal tip short, Right cheliped larger in male, left cheliped larger in stout, with a slight bend; membrane commences ovigerous female. Upper surface of merus granular shortly behind the tip; inner row of 4-5 bristles very with two well-developed spines and five to six sparsely placed commencing distantly from distal tip granules on its anterior margin; no spine on poster- and terminate before outer row of bristles. Female ior margin. Carpus with tuberculated upper surface gonopore slightly reniform and located on thoracic bears one large spine on its inner angle and three sternite 6, posterior to suture 5/6 and corresponds to small spines on outer angle. Manus tumid, five the bases of second pair of pereiopods. costate, with five spines on its upper surface; two blunt spines on distal margin immediately posterior to articulation with dactylus; two well-developed Description spines in the middle of upper surface trailed by coarse granular costae; fifth spine on the posterior Carapace (CW It =26.68 + 4.44 mm; CL margin just anterior to articulation with carpus; 15.85 +3.94 mm; CW/CL u =1.71 +0.16) evenly outer surface with three smooth costae, the lower- tomentose/pilose, with the exception of transverse most reaches the tip of the dactylus. Dactylus ridges and swellings (Figure 3A,B). Frontal margin smooth, strong, and slender with curved tips, longer cut into six teeth (excluding inner supra-orbital than manus. angles), median frontal teeth bluntly triangular, Ambulatory legs long, slender; third pair longest, projecting beyond broadly rounded sub-medians, although shorter than the chelipeds. Dactyli and laterals triangular, narrowest, separated from sub- propodi of first three pairs of legs covered with medians by wide, U-shaped notch (Figure 3C). densely packed bristles on both anterior and poster- Orbits (OD is = 3.05 +0.56 mm) with strong dorsal ior margins, carpus and merus bear bristles only on inclination; major orbital diameter two-fifths to posterior margin. Posterior margin of propodus and nearly half of the inter-orbital space (IS tt =6.26 ± carpus of natatory leg smooth, merus with promi- 1.52 mm; OD/IS s =0.49 +0.03). Inter-orbital nent spine on its postero-distal margin. space almost one-quarter the carapace width (IS/ Thoracic sternites and abdominal cavity in male CW IA =0.23 +o.on). Fronto-orbital margin (FW sternum finely granular (Figure 3F). First three u =12.77 +2.85 mm) almost half the carapace thoracic sternites narrow, sutures 1/2 and 2/3 width; antero-lateral margins oblique and more or prominent. Suture 3/4 wide and shallow. Sternite 4 less arched. Infra-orbital lobe with row of granules widest, medially divided by a shallow anterior on frontal margin. Antennae fold transversely. Basal extension of abdominal cavity. Abdominal cavity antenna! joint • with smooth granular ridge, its commences from posterior portion of sternite 4 prolongation in contact with front, thus excluding and covers sternites 5-8. Deep groove along abdom- antennal flagellum from orbit. Merus of third max- inal cavity medially divides sternites 5-8. Granule of illiped with outer distal angle produced sideways. abdominal lock located immediately anterior to Antero-lateral margin of carapace moderately suture 5/6. Thoracic sternites in female specimens arched, cut into six serrated teeth separated by wide similar to those in male (Figure 3G). notches (Figure 3D). First tooth slightly notched, Sixth abdominal segment in males broader than second and third sub-equal, blunt pointed with long, its sides parallel over less than half their lengths serrated margins, fourth and fifth wider and spini- and then gradually converge distally (Figure 3H); form, sixth tooth greater than twice the length of the basal margin slightly convex, distal margin concave. preceding five and laterally directed (Figure 3D). GI L-shaped, its distal tip short, stout, with a slight New species of portunid crab 583 •

10 mm

C

(Iv)

G

J

1--1 1---I 1mm 1mm 1mm Figure 3 (Continued) 584 V. P Padate et al. bend (Figure 4A), reaches sixth thoracic sternite; Habit membrane commences shortly behind the tip; outer Charybdis goaensis was mainly collected from a trawl row of bristles commences close to distal tip and operation in the estuary. However, capture of an terminates in the neck region; bristles clumped ovigerous female specimen in an offshore trawl from towards the distal end and sparsely placed towards a depth of 13-14 m in March 2007 indicates the the neck; inner row of 4-5 bristles very sparsely probability of catadromous habit of the females of placed commencing distantly from distal tip and this species for spawning. terminate before outer row of bristles; bristles on abdominal margin randomly and very widely placed (Figure 4B,C). Remarks Females exhibit dimorphism in abdominal shape; Crabs were identified up to the sub-genus level using immature females with narrow, triangular abdomen orthodox taxonomic studies based on morphological (Figure 31), wide watch glass-shaped abdomen in characters described by Alcock (1899), Leene ovigerous female (Figure 3J). Female gonopore (1938), Sakai (1976) and Wee & Ng (1995). The slightly reniform, located on thoracic sternite 6, distinguishing morphological characteristics of the posterior to suture 5/6 and corresponding to the sub-genus Charybdis are: lobule at external angle of bases of second pair of pereiopods (Figure 3G). basal antennal joint in contact with front and completely excludes flagellum from orbital hiatus; Colour posterior angles of carapace may be accented or not, but posterior margin forms a curve with postero- Carapace light brown over tomentose regions, po- lateral margins; four median frontal teeth not lished surfaces of granular transverse ridges and dissimilar to lateral frontal teeth; antero-lateral regional tuberosities dark brown; pale brown to off- margins cut into six teeth (or seven, in which case white ventrally. Chelipeds dark brown with whitish there are six large and one small teeth); no spine on tips; other pereiopods light brown. posterior margin of cheliped (Leene 1938; Sakai 1976). Etymology The distinctiveness of the new species was ascer- The species name,. Charybdis (Charybdis) goaensis is tained based on the comparison with morphological derived from the type locality, Goa, west coast of descriptions of its congeners (Fabricius 1787; Miers India. 1884; Alcock 1899; Rathbun 1923a, 1923b, 1924; Shen 1934; Leene 1938; Ward 1941; Leene & Buitendijk 1949; Edmondson 1954; Rees & Distribution Stephenson 1966; Stephenson & Rees 1967b, Charybdis goaensis is currently known only from the 1968; Zarenkov 1970; Sakai 1976; Crosnier 1984; type locality (Zuari Estuary) and offshore fishing Wee & Ng 1995; Spiridonov & Tiirkay 2001). grounds of Goa, west coast of India. Morphological characters such as number of antero-lateral teeth, presence of transverse ridges on carapace, armature of chelipeds and natatory legs Habitat characteristics were used as criteria to differentiate the new species The specimens were collected from 6 to 7 m depth from its congeners. Based on the above criteria, C. localities from the Zuari Estuary, Goa, west coast of goaensis with six antero-lateral teeth differed with C. India during September (end of the south-west heterodon and C. demani with five and seven antero- monsoon). At the time of collection, salinity re- lateral teeth, respectively. The new species differed corded was in the range of 22-32 ppt. The collection with 12 other species (C. rathbuni, C. salehensis, C. site is in the close vicinity of Mormugao Port Trust, variegata, C. brevispinosa, C. granulata, C. natator, and the waterway is subjected to regular dredging. C. seychellensis, C. beuuforti, C. curtilobus, C. cookei,

Figure 3. Charybdis (Charybdis) goaensis sp. nov. A, Dorsal surface of carapace (camera lucida diagram). B, Dorsal surface of carapace (photograph). C, Frontal margin of carapace (camera lucida diagram). D, Antero-lateral margin of carapace (camera lucida diagram). E, Dorsal view of right cheliped (camera lucida diagram). F, Ventral surface indicating thoracic sternites and abdominal cavity of male (camera lucida diagram). (3, Ventral surface indicating thoracic sternites and abdominal cavity of female (camera lucida diagram). H, Male abdomen (camera lucida diagram). 1, Abdomen of immature female (camera lucida diagram). J, Abdomen of mature female (camera lucida diagram). (i) Swellings on carapace devoid of tomentum; (ii) tomentose carapace; (iii) wide U-shaped notch separating sub-median and lateral frontal teeth; (iv) bluntly triangular median frontal teeth; (v) first untero-lateral tooth slightly notched; (vi) wide notches separate the antero-lateral teeth; (vii) last antero-lateral tooth largest and laterally directed; (viii) 2 spines with blunt tips on upper surface of manus; (ix) 3 well-developed spines on upper surface of manus; (x) 2 spines on anterior margin of merus; (xi) grain of abdominal lock (pair); (xii) thoracic sternites 1-3; (xiii) thoracic sternites 4-8; (xiv) abdominal cavity; (xv) female gonopore (pair); (xvi) sixth abdominal segment. New species of portunid crab 585 A and 'distinctly bifid first antero-lateral tooth' and `three spines on cheliped manus' in C. ihlei. The identification of the new species was further confirmed through thorough comparison of its mor- phological characteristics with those of the holotype (AMNH 8382) and male paratype (AMNH 7893) specimens of its closest congener, C. philippinensis, obtained from the American Museum of Natural History. A brief updated description of C. •philippi- B nensis along with its morphometry and diagnostic morphological illustrations (camera lucida diagrams and photographs) is provided below.

Charybdis (Charybdis) philippinensis Ward, 1941 Comparison of description of Ward and diagnosis of holotype specimen (Catalogue No. AMNH 8382 obtained from American Museum of Natural History) (The holotype specimen obtained was devoid of the right cheliped and legs on right side)

Carapace (CW µ = 35.65 ±2.56 mm; CLµ = 20.50 ± 1.30 mm; CW/CL u =1.74 ±0.01 mm) naked, de- void of tomentum (Figure 5A). Median frontal teeth rounded, sub-median and lateral frontal teeth sepa- rated by narrow V-shaped notch (Figure 5C). Orbits Figure 4. Charybdis (Charybdis) goaensis sp. nov. Scanning elec- tron micrographs (SEM) of first pleopod or gonopod (GI) (left). with strong dorsal inclination; major orbital diameter (a) Entire gonopod. (b) Tip of gonopod. (c) Tip of gonopod (OD 1.1 =3.97 + 0.18 mm) almost half the length of (enlarged view). the inter-orbital space (IS IA =8.46 +0.45 mm; OD/ IS = 0.47 + 0.00) (Figure 5A). Inter-orbital space almost one-quarter the carapace width (IS/CW p = C. rostrata and C. callianassa) by the absence of 0.24). Antero-lateral teeth six in number, first slightly transverse ridges posterior to last antero-lateral notched, sixth greater than two times the length of the teeth. Further, C. goaensis with two spines on preceding five and laterally directed; teeth separated anterior margin of merus of cheliped differed with by narrow notches (Figure 5D). Transverse ridges of 24 other species (C. orientalis, C. hawaiensis, C. carapace faintly granular, two proto-gastric, one feriata, C. padadiana, C. rosaea, C. annulata, C. meso-gastric, a medially divided meta-gastric, and hellerii, C. vannamei, C. spinifera, C. acuta, C. incisa, epibranchials along the level of the last antero-lateral C. javanensis, C. lucifera, C. amboinensis, C. yaldwyni, teeth; no transverse ridges beyond the level of the last C. jaubertensis, C. japonica, C. affinis, C. acutidens, C. antero-lateral teeth (Figure 5A). Cardiac region rufodactylus, C. sagamiensis, C. riversandersoni, C. devoid of tuberosity (Figure 5A). Chelipeds sub- crosnieri, C. holosericus) with three spines, and two equal (LC p =36.30 + 3.96 mm; RC p =40.08 +2.15 species (C. miles, C. meteor) with four to five spines, mm), smooth; merus with two well-developed spines respectively. and five to six granules on its anterior margin; mantis C. goaensis was observed to be morphologically tumid, five-costate, with five spines on upper.surface, most similar to C. philippinensis, C. ihlei and C. two blunt spines on distal margin immediately poster- anisodon based on 'two spines on anterior margin of ior to articulation with dactylus; dactylus smooth, merus of cheliped', and 'absence of spines on strong, and slender with curved tips, longer than posterior margin of propodus of natatory leg'. manus (Figure 5B). Propodus of natatory legs Further, bdth C. goaensis and C. philippinensis smooth, merus bears prominent spine on its pos- possess 'slightly notched first antero-lateral tooth' tero-distal margin. Penultimate abdominal segment and 'five spines on cheliped manus' and differed broader than long, its sides parallel over less than half from 'obtusely triangular first antero-lateral tooth' their lengths and then gradually converge distally and 'two spines on cheliped manus' in C. anisodon, (Figure 5B).

586 V P Padate et al.

B

10 mm

(Iv)

C

(v)

Figure 5. Charybdis (Charybdis) philippinensis Ward, 1941. A, Dorsal surface of carapace (photograph). B, Ventral surface of carapace (photograph). C, Frontal margin of carapace (camera lucida diagram). D, Antero-lateral margin of carapace (camera lucida diagram). (i) Narrow V-shaped notch separating sub-median and lateral frontal teeth; (ii) rounded median frontal teeth; (iii) first antero-lateral tooth slightly notched; (iv) narrow notches separate the antero-lateral teeth; (v) last antero-lateral tooth largest and laterally directed.

Comparison between the new species and its 2. Antero-lateral margins of carapace appear to be closest congener, C. philippinensis, revealed that the divided into five teeth as first two teeth are new species possessed 'bluntly triangular frontal grown together except for distal end rendering teeth; wide U-shaped notch separating sub-median `bifurcated first tooth' appearance . . . Charybdis and lateral frontal teeth' as compared to 'rounded (Charybdis) heterodon Nobili, 1906. median frontal teeth; narrow V-shaped notch separ- —Antero-lateral margins of carapace appear to be ating sub-median and lateral frontal teeth' of the divided into seven teeth due to division of latter species. In addition, antero-lateral teeth of first tooth into larger (anterior) and smaller C. goaensis were separated by 'wide notches' as (posterior) teeth Charybdis (Charybdis) compared to 'narrow' ones of C. philippinensis. demani Leene, 1937. Moreover, C. goaensis differed from the latter species - Antero-lateral margins of carapace divided into by the presence of 'granular tuberosity' on cardiac six teeth 3 region. 3. Two spines on the anterior margin of merus of A taxonomic key to 43 species of the subgenus cheliped. First antero-lateral tooth square-cut to Charybdis is provided below to substantiate our obtusely triangular; last tooth greater than twice claim of identification of a new portunid crab species in length than any of the preceding teeth, and from Goa, west coast of India. projected laterally. Posterior margin of propodus of natatory (last) leg smooth 4 —Three spines on the anterior margin of merus of Key to the species of swimming crabs of the cheliped. First antero-lateral tooth usually smal- sub-genus Charybdis ler than succeeding ones, may be truncated; 1. Transverse ridges on carapace posterior to the length of last antero-lateral tooth may or may not last antero-lateral tooth absent 2 exceed that of preceding spine, may be projected — Transverse ridges on carapace posterior to the laterally. Posterior margin of propodus of nata- last antero-latetal tooth present 28. tory leg serrated or smooth New species of portunid crab 587 - Three to five spines on the anterior margin of - All five spines on upper surface of manus of merus of cheliped. First antero-lateral tooth cheliped well developed 12. smallest, may or may not be truncated, last 11. Margin of median frontal teeth well in one largest and either forwardly, laterally or advance of that of sub-medians; presence of backwardly directed. Posterior margin of pro- faint transverse ridges on frontal region of podus of natatory leg serrated 22. carapace Charybdis (Charybdis) rosaea 4. First antero-lateral tooth obtusely triangular. (Hombron & Jacquinot, 1846). Two spines on upper surface of manus - Margin of median frontal teeth slightly in of cheliped Charybdis (Charybdis) advance of that of sub-medians; no transverse anisodon (De Haan, 1833). ridge on frontal region Charybdis First antero-lateral tooth distinctly bifid. Three (Charybdis) annulata (Fabricius, 1798). spines on upper surface of manus of cheli- 12. Posterior margin of carpus of natatory leg bears ped Charybdis (Charybdis) ihlei Leene & spine 13. Buitendijk, 1949. Posterior margin of carpus of natatory leg First antero-lateral tooth slightly notched. smooth, lacks spine ,, 15. Five spines on upper surface of manus of 13. Last antero-lateral spine elongate, projects be- cheliped 5 yond preceding spine 14. 5. Median frontal teeth bluntly triangular, sub- - Last antero-lateral spine smallest, does not medians and laterals separated by wide U- project beyond preceding spineCharybdis (Char- shaped notch. Antero-lateral teeth separated by ybdis) vannamei Ward, 1941. wide notches. Prominent granular tuberosity on 14. Median frontal teeth sharp, triangular, project- cardiac region Charybdis (Charybdis) ing beyond narrow sub-medians; posterior goaensis sp. nov. margin of propodus of natatory leg denticula- - Median frontal teeth rounded, sub-medians ted Charybdis (Charybdis) hellerii and laterals separated by narrow V-shaped (A. Milne Edwards, 1867). notch. Antero-lateral teeth separated by narrow Median and sub-median frontal teeth broadly notches. Lack of tuberosity on cardiac re- rounded, medians project slightly in advance of gion Charybdis (Charybdis) philippinensis sub-medians; posterior margin of propodus of Ward, 1941. natatory leg smooth Charybdis 6. Upper surface of manus of cheliped with four (Charybdis) spinifera (Miers, 1884). spines 7 15. Posterior margin of propodus of natatory leg Upper surface of manus of cheliped with five serrated 16. spines 10. - Posterior margin of propodus of natatory leg 7. First antero-lateral tooth spiniform, second smooth 21. rudimentary, attached to posterior margin of 16. All frontal teeth sharply triangular. . . . Charybdis first 8 (Charybdis) acuta (A. Milne Edwards, 1869). First antero-lateral tooth notched or blunt, - All frontal teeth rounded 17. second not rudimentary 9 - Frontal teeth either bluntly triangular or 8. Inner surface of manus of cheliped smooth; rounded 18. merus of fifth leg proportionally longer 17. Median frontal teeth slightly in advance of sub- Charybdis (Charybdis) orientalis Dana, medians, laterals least advanced . . . . Charybdis 1852. (Charybdis) incisa Rathbun, 1923a. Inner surface of manus of cheliped strongly - Median frontal teeth well in advance of granular; merus of fifth leg relatively shorter sub-medians, laterals slightly in advance Charybdis (Charybdis) hawaiensis of sub-medians Charybdis (Charybdis) Edmondson, 1954. javanensis Zarenkov, 1970.. 9. First antero-lateral tooth distinctly bifid, second 18. Frontal teeth bluntly triangular; median and fifth teeth larger than first, last tooth teeth blunter than sub-medians and produced laterally Charybdis (Charybdis) laterals 19. .feriata (Linnaeus, 1758) Median and sub-median frontal teeth broadly - First two antero-lateral teeth blunt, second and rounded, laterals broadly triangular 20. fifth teeth smallest, last tooth not produced 19. Sub-median frontal teeth separated from laterally . . . . ,Charybdis (Charybdis) padadiana the laterals by deep V-shaped incision. Trans- Ward, 1941. verse ridges of carapace faintly granu- 10. Manus of cheliped bears five spines on upper sur- lar Charybdis (Charybdis) lucifera face, two on distal margin tuberculated 11. (Fabricius, 1798). 588 V P Padate et al. - Sub-median frontal teeth separated from the styliform; manus slender, devoid of squamiform laterals by wide and deep fissure. Transverse marking on outer side 26. ridges of carapace distinctly granular 25. Carina on inner as well as outer surface of Charybdis (Charybdis) manus in the form of conspicuous ridges; amboinensis Leene, 1938. movable finger long, slender and deeply 20. Cardiac region of carapace with corrugated grooved; postero-distal margin of propodus of surface; first antero-lateral tooth markedly trun- natatory leg smooth . . . Charybdis (Charybdis) cated; outer surface of manus strongly squami- rufodactylus Stephenson & Rees, 1968.. form throughout; G1 of male short, stout and - Carina on inner as well as outer surface of sharply curved, absence of upper inner spines manus in the form of rounded ridges; movable proximal to lip of G1 Chary- finger short, relatively stout and shallow groo- bdis (Charybdis) yaldwyni Rees & Stephenson, ved; two spinules on postero-distal margin of 1966. propodus of natatory leg Charybdis - Cardiac region of carapace typically granulated; (Charybdis) sagamiensis Parisi, 1916. first antero-lateral tooth slightly truncated; 26. Inner supra-orbital lobe styliform, outer outer surface of manus coarsely granular on sharpened; dactyli of second to fourth pereiopods upper half and squamiform on lower half; GI thin, curved with lancet-shaped tip . . . Charybdis of male long, slender with sinuous margin, (Charybdis) acutidens Tiirkay, 1986. presence of inconspicuous row of well-spaced - Inner supra-orbital lobe broadly to sharply spinules proximal to lip Charybdis triangular; dactyli of second to fourth pereio- (Charybdis) jaubertensis Rathbun, 1924. pods moderately wide, slightly curved. . . . 27. 21. Manus of cheliped with well developed spines; 27. Median frontal teeth acutely rounded; lateral penultimate segment of male abdomen with frontals with outer edge convex; basal antennal convex lateral borders; last antero-lateral tooth joint bearing distinct granules; lip of G1 of male smallest and spiniform, not projecting beyond tapering with conspicuous spinules evenly distrib- preceding tooth Charybdis (Charybdis) uted along edges; lateral spines not closely set, japonica (A. Milne Edwards, 1861). extending to about one-fifth of the neck - Manus of cheliped with poorly developed Charybdis (Charybdis) riversandersoni Alcock, spines; penultimate segment of male abdomen 1899. with lateral margins parallel for proximal half; - Median frontal teeth sharply granular; lateral last antero-lateral tooth elongated, projecting frontals with straight edges; basal antennal joint laterally beyond preceding tooth . . . Charybdis bearing fused granules; lip of G1 of male elon- (Charybdis) affinis Dana, 1852. gated and bent downward, spinules concentrated 22. Carapace pilose and narrow, CW/CL ratio on abdominal edge near tip; lateral spines being 1.41, ridges of carapace distinct; lower closely set, extending to one-quarter of the neck surface of manus with squamiform mar- Charybdis (Charybdis) crosnieri king Charybdis (Charybdis) miles Spiridonov & Turkay, 2001. (De Haan, 1835). 28. Transverse ridge on cardiac region interrupted - Carapace generally lacking pilosity, relatively 29. broad, CW/CL ratio being 1.49-1.67, ridges of - Transverse ridge on cardiac region not inter- carapace faint; squamiform marking on manus, rupted 36. if present may be faint 23. 29. Small-sized species; mesobranchial region 23. Ridges on dorsal surface of carapace finely of carapace with two pairs of transverse ridges granular, mesogastric ridges distinct; third to 30. fifth antero-la'teral teeth with straight edges, - Species of a good size; mesobranchial region of others with sinuous edges Charybdis carapace with two to three pairs of transverse (Charybdis) meteor Spiridonov & Tiirkay, 2001. ridges 33. - Ridges on carapace finely or distinctly 30. Second antero-lateral tooth of carapace granular; all antero-lateral teeth with sinuous smaller than the first, and attached to its edges 24. posterior margin Charybdis (Charybdis) 24. Transverse ridges of carapace distinctly granu- rathbuni Leene? 1938. lar; inner sub-orbital lobe broadly triangular; - Second antero-lateral tooth as large as the first, manus massive with faint squamiform marking not attached to the posterior margin of the on lower surface of outer side 25. latter 31. Transverse ridges of carapace finely granular; 31. Sixth antero-lateral tooth largest, spiniform and inner sub-orbital lobe sharply triangular or laterally projected 32. New species of portunid crab 589

- Sixth antero-lateral tooth distinctly smaller than Conclusion Charybdis preceding tooth, laterally projected. . . . In view of the above description and comparison Leene, 1938. (Charybdis) salehensis of morphological characters with its congeners, 32. All frontal teeth acuminate at tip. Sub-distal Charybdis goaensis is a new species. The distinguish- portion of male gonopod separated as distinct ing characters of the new species are as follows. lobe from tip Charybdis (Charybdis) variegata (Fabricius, 1798). 1. 'Median frontal teeth bluntly triangular'. - Only lateral frontal teeth acuminate at tip. Sub- 2. 'Sub-median and lateral frontal teeth separated distal portion of male gonopod not distinct from by wide, U-shaped notch'. tip Charybdis (Charybdis) brevispinosa 3. 'First antero-lateral tooth slightly notched'. Leene, 1937 4. 'Last tooth of antero-lateral margin greater 33. Dorsal surface of carapace even and uniformly than twice the length of the preceding five covered with soft tomentum. Penultimate seg- and laterally directed'. ment of male abdomen not convex on lateral 5. `Antero-lateral teeth separated by wide margins 34. notches'. - Dorsal surface of carapace somewhat uneven, 6. 'Prominent granular tuberosity on cardiac unevenly tomentose. Penultimate segment of region'. male abdomen convex on lateral margins 7. 'Two spines on the anterior margin of merus of Charybdis (Charybdis) granulata (De cheliped'. Haan, 1833). 8. 'Five spines on upper surface of manus of 34. First antero-lateral tooth of carapace trun- cheliped, two on distal margin remain tubercu- cated at tip. Frontal ridge of carapace lated'. absent Charybdis (Charybdis) natator 9. 'Membrane on tip of G1 of male commences (Herbst, 1789). shortly behind the tip; row of sparsely-placed — First antero-lateral tooth not truncated. Frontal bristles on outer margin starts at some distance ridge present 35. from the tip, but anterior to the above mem- 35. Carapace with high relief and prominent brane; row of 4-5 bristles very sparsely placed transverse granular ridges. Granules on on inner margin commencing distantly from chelipeds very prominent and projecting distal tip and terminating prior to outer row of Charybdis (Charybdis) beauforti bristles; bristles on abdominal margin ran- Leene & Bnitendijk, 1949. domly and very widely placed'. Carapace with low relief and less prominent transverse granular ridges. Granules on chelipeds less prominent Charybdis Acknowledgements (Charybdis) seychellensis Crosnier, 1984. The authors are grateful to the Ballast Water 36. Mesobranchial region of carapace with two pairs Management Programme, India executed by Na- of transverse ridges. Two spines on anterior tional Institute of Oceanography, Dona Paula, Goa margin of merus of cheliped Charybdis for Directorate General of Shipping, Ministry of (Charybdis) curtilobus (Stephenson & Rees, Shipping, Government of India. We would also take 1967b). this opportunity to express our deep gratitude to - Mesobranchial region of carapace with one pair Dr B.F. Chhapgar, an eminent carcinologist from of transverse ridges. Three spines on anterior the Indian Ocean region for his valuable guidance margin of merus of cheliped Charybdis in confirming the identity of the newly described (Charybdis) cookei Rathbun, 1923b. species. We gratefully acknowledge Dr Sara - No transverse ridges on mesobranchial re- C. Watson and Dr Mark Siddall, American gion 37. Museum of Natural History, for their timely help 37. Carapace about four-fifths as long as broad; two in providing Charybdis philippinensis specimens on spines on palm of cheliped. Lateral margins loan. We are also indebted to Mr V. Khedekar, of sixth abdominal segment of male curved Geological Oceanography Division, NIO for the Charybdis (Charybdis) rostrata Scanning Electron Microscope (SEM) photography (A. Milne Edwards, 1861). of the crab specimens. The authors are extremely - Carapace about two-thirds as long as broad; grateful to the anonymous reviewers for their three spines on palm of cheliped. Lateral critical suggestion which enabled improvement of margins of sixth abdominal segment of male the manuscript. The help extended by Dr M.P. parallel for half their lengths Charybdis Tapaswi in obtaining the references from different (Charybdis) callianasa (Herbst, 1801). places without which this study would have been 590 V P Padate et al.

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