Biota of India - Subba Rao

AQUATIC CONSERVATION: MARINE AND FRESHWATER ECOSYSTEMS
Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Published online 14 January 2005 in Wiley InterScience
(www.interscience.wiley.com). DOI: 10.1002/aqc.659
Comprehensive review of the records of the biota of the

Indian Seas and introduction of non-indigenous species
D.V. SUBBA RAO*

HED, Marine Environmental Sciences Division, Bedford Institute of Oceanography,
Dartmouth, Nova Scotia, Canada
ABSTRACT
1. Comparison of the pre-1960 faunal survey data for the Indian Seas with that for the post-1960
period showed that 205 non-indigenous taxa were introduced in the post-1960 period; shipping
activity is considered a plausible major vector for many of these introductions.
2. Of the non-indigenous taxa, 21% were fish, followed by Polychaeta (511%), Algae (10%),
Crustacea (10%), Mollusca (10%), Ciliata (8%), Fungi (7%), Ascidians (6%) and minor
invertebrates (17%).
3. An analysis of the data suggests a correspondence between the shipping routes between India and
various regions. There were 75 species common to the Indian Seas and the coastal seas of China and
Japan, 63 to the Indo-Malaysian region, 42 to the Mediterranean, 40 and 34 to western and eastern
Atlantic respectively, and 41 to Australia and New Zealand. A further 33 species were common to the
Caribbean region, 32 to the eastern Pacific, 14 and 24 to the west and east coasts of Africa respectively,
18 to the Baltic, 15 to the middle Arabian Gulf and Red Sea, and 10 to the Brazilian coast.
4. The Indo-Malaysian region can be identified as a centre of xenodiversity for biota from
Southeast Asia, China, Japan, Philippines and Australian regions.
5. Of the introduced species, the bivalve Mytilopsis sallei and the serpulid Ficopomatus enigmaticus
have become pests in the Indian Seas, consistent with the Williamson and Fitter ‘tens rule’. Included
amongst the biota with economic impact are nine fouling and six wood-destroying organisms.
6. Novel occurrences of the human pathogenic vibrios, e.g. Vibrio parahaemolyticus, non-01 Vibrio
cholerae, Vibrio vulnificus and Vibrio mimicus and the harmful algal bloom species Alexandrium spp.
and Gymnodinium nagasakiense in the Indian coastal waters could be attributed to ballast water
introductions.
7. Introductions of alien biota could pose a threat to the highly productive tropical coastal waters,
estuaries and mariculture sites and could cause economic impacts and ecological surprises.
8. In addition to strict enforcement of a national quarantine policy on ballast water discharges,
long-term multidisciplinary research on ballast water invaders is crucial to enhance our
understanding of the biodiversity and functioning of the ecosystem.
Copyright # 2005 John Wiley & Sons, Ltd.
KEY WORDS: ballast water introductions; biota of the Indian Seas; non-indigenous species; introduced species;
biological invasions
*Correspondence to: D.V. Subba Rao, HED, Marine Environmental Sciences Division, Bedford Institute of Oceanography,
PO Box 1006, Dartmouth, Nova Scotia B2Y 4A2, Canada. E-mail: durvasulaSR@mar.dfo-mpo.gc.ca
Copyright # 2005 John Wiley & Sons, Ltd. Received 29 April 2003
Accepted 7 July 2004
118 D.V. SUBBA RAO
INTRODUCTION
Since 1878, faunal surveys have been carried out in the waters around peninsular India, viz. Bay of Bengal,
Arabian Sea, and Andaman Sea, collectively referred to as the Indian Seas, through oceanographic
expeditions and by British surgeons with a flair for natural history, universities and government
laboratories. These studies resulted in a vast body of data collected between 1865 and 1960, providing a
solid baseline for what is accepted as the natural biota of the Indian Seas. In 1961, the Shipping
Corporation of India was formed, with the merger of Eastern Shipping and Western Shipping. This
signalled an increase in India’s modern shipping activity, and is the reason for the choice of 1960 as the
baseline. Against this background, data collected from 111 papers published between 1960 and 2004 reveal
205 taxa never before reported as present in the Indian Seas region. These represent 32 taxonomic
groups, ranging from fungi to fish, that have been introduced into the Indian Seas from their native
waters. This paper describes the physical area of interest (the Indian Seas), the baseline data (1878–1960),
the data relating to the new taxa, and the probable mechanisms of the introductions. In addition, the
probable consequences of the introductions and the action that might be used to combat them are
discussed.
The geographic area

The Indian Seas cover 3 106 km2, an area considerably larger than the land mass of India. Their total
shelf area of about 0.84 106 km2 approximates that of the Gulf of Mexico plus the Caribbean Sea. The
Indian coastline is 7515 km in total length, with 12 major ports and 40 minor ports (Figure 1). The Bay of
Bengal and the Arabian Sea contribute 59% and 41% respectively to the 2.02 106 km2 Indian Exclusive
Economic Zone (EEZ). These waters contain distinct habitats, such as estuaries, mangrove swamps,
brackish water lakes, coral reefs, islands and offshore waters, and support a great diversity of fauna and
flora. Additionally, brackish water lakes, such as the Chilika on the east coast of India, and several
mariculture operations produce a rich commercial harvest. The total estimated fishery potential for the
Indian EEZ is 3.92 106 t, compared with the 2.69 106 t currently harvested, of which 60% is from the
Arabian Sea (Somvanshi, 1998). More than 20% of India’s population is coastal dwelling, which greatly
increases the burden on the coastal zone through urban settlements, fishing, mariculture, marine transport,
recreation, offshore oil and gas exploration, and dumping of wastes.
Baseline data on biota of the Indian Seas

Our knowledge of the fauna and flora in the Indian Seas between 1874 and 1960 is based on descriptive
surveys around India. Hundreds of taxa, old and new, were described in about 150 publications, and the
results serve as the baseline for indigenous fauna and flora of the seas around India.
Publications on distribution of marine fauna and flora

It is evident from the number of publications on marine sciences in the Indian Seas that a good knowledge
base of the fauna and flora of the area exists (Table 1). References to the literature on the fauna and flora of
the Indian Seas can be found in a number of extensive bibliographies (see Yentsch, 1962; Alagaraswami
et al., 1968; Sriramachandra Murty et al., 1968, 1969; Jones, 1971; Zeitzschel, 1973; Santhakumaran, 1985;
Subba Rao, 2002).
There has been a great increase in the number of publications in marine sciences since 1961, suggesting
greater awareness of these sciences (Table 1). Of these, emphasis is in two areas, namely the
marine resources (such as the fish, fisheries and the biota) and the monsoons, because of their impact on
droughts and floods and on Indian agriculture. Several young Indian scientists with a penchant for
Copyright # 2005 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)
BIOTA OF THE INDIAN SEAS 119
Figure 1. Indian Seas with major ports. Broken line demarcates the EEZ in the Indian Seas.
Table 1. Number of abstracts on Indian Ocean region in Aquatic Sciences and Fisheries cited by
Cambridge Scientific Abstracts and Oceanic Abstracts
Category of study Up to 1960 1961 to 2004

Fish, fishery, fisheries 910 7717
Organisms other than fish and fisheries 1346 5024
Geological oceanography 24 422
Physical oceanography 313 523
Chemical oceanography 24 245
Monsoons 196 1647
120 D.V. SUBBA RAO
Table 2. List of groups that have been studied in detail in the Indian Seas
Number of taxa
reported Reference
Taxonomic group
Fungi 88 Prasannarai et al. (1999), Prasannarai and Sridhar (2001)
Diatoms 176 Subrahmanyan (1946)
Dinoflagellates 108 Subrahmanyan (1968, 1971)
Foraminifera 85 Kameswara Rao and Satyanarayana Rao (1979)
Tintinnids 47 Krishnamurti et al. (1979)
Porifera 481 Thomas (1983)
Hydromedusae 80 Santhakumari and Nair (1998)
Siphonophora 37 Rengarajan (1973, 1983)
Scleratinia corals 199 Gopinadha Pillai (1983)
Polychaetes 120 Srikrishnadhas et al. (1987)
Ectoproct bryozoa 22 Satyanarayana Rao (1992)
Chaetognatha 19 Nair and Rao (1973)
Ostracoda 40 Annapurna and Sarma (1986)
Cumacea 242 Kurian (1965)
Cirriped barnacles 15 Karande (1965)
Copepoda 400 Sewell (1912a,b,1929,1947,1948)
Amphipoda } Gammaridae 78 Nagappan Nayar (1965)
Pelagic Amphipoda 86 Krishna Pillai (1965a,b)
Sphaeromidae 5 Krishna Pillai (1965c)
Mysidacea 75 Krishna Pillai (1965d)
Brachyura 208 Laurie (1906)
Stomatopoda 18 Alikunhi (1965)
Euphausiacea 23 Sebastian (1965)
Limnoridae 7 Santhakumaran and Srinivasan (1988)
Echinodermata 200 James (1983)
Teredinidae 34 Santhakumaran (1985)
Cephalopoda 27 Silas et al. (1985a,b)
Heteropoda 23 Richter (1974)
Thecosomatous pteropoda 22 Richter (1979)
Appendicularians 26 Fenaux (1973)
Ascidians 28 Renganathan and Krishnaswamy (1985)
85 Meenakshi (1998)
Fish 1400 Talwar and Kacker (1984)
Commercial fish 548 Talwar and Kacker (1984)
Habitat Group
Intertidal macro algae 80 Rao MU (1970, 1999, 2001)
Interstitial sand-dwelling fauna 199 Chandrasekhara Rao (1975)
Coral reef fauna 199 Gopinadha Pillai (1983)
Bottom fauna 208 Parulekar et al. (1981)
Fouling organisms 54 Ganapati et al. (1958)
Meiofauna 65 Sarma and Ganapati (1974)
Mangroves 58 Blasco (1977)
Mangrove-associated biota 546 Kathiresan (2000)
taxonomy have made significant contributions to a large body of data that facilitates comparison with
baseline data on the distribution of the biota for this region. A great diversity of marine fauna and
flora exists in the Indian Seas (Table 2), with most of the major taxonomic groups represented by a large
number of taxa.
NON-NATIVE TAXA
Even today, reports of occurrences of taxa not native to the Indian Seas have been in the form of very brief
notes and mainly confined to obscure journals with limited circulations. Consequently, they have not
received much scientific attention. For this analysis, records of genera and species new to science are not
included; only those reports of species found for the first time in Indian waters, the Arabian Sea and the
Bay of Bengal have been reviewed. The waters around the Laccadives and Maldives are considered part of
the Arabian Sea, and those surrounding the Andamans, Nicobars and Sri Lanka are a part of the Bay of
Bengal. However, the seas around the islands have remained inadequately sampled, and the few ‘new’
records of taxa have turned out to be first time records to these islands, but not so for the Bay of Bengal or
the Arabian Sea. Additionally, there has historically been regular intra-coastal shipping activity between
these island waters and the Bay of Bengal and the Arabian Sea that would have facilitated shuffling of the
biota within their natural range (Eno, 1996).
Collation of details on non-native taxa

Of the thousands of publications on the coastal seas around India, 111 publications described biota new to
the region (Table 3). During 1960–1969 there were only five publications, compared with a peak of 64
during the next decade, in which there were 115 taxa reported new to the region. The taxa consisted of 32
major taxonomic groups (Table 4 and Appendix). The bulk of the organisms introduced were fish (21%),
Polychaeta (11%), Algae (10%), Crustacea (10%), Mollusca (10%), Ciliata (8%), Fungi (7%), Ascidians
(6%) and others (17%).
This review shows that 205 taxa (see Appendix), not native to the Indian waters, have been introduced
since 1960 from various seas (Figure 2). For example 75 species were common to the coastal seas of China
and Japan, 63 to the Indo-Malaysian region, 42 to the Mediterranean, 40 to the western Atlantic, and 41 to
Australia and New Zealand. Next in importance are 34 taxa from the eastern Atlantic, 33 from the
Carribean region, and 29 from the eastern Pacific. Contributions of other regions were low, with 24 from
East Africa, 18 from the Baltic, 15 from the middle Arabian Gulf and Red Sea, 14 from the west coast of
Africa and 10 from the Brazilian coast (Figure 2).
Several non-indigenous fouling and wood-destroying organisms that are of economic importance
were also present (see Appendix). Besides one diatom, nine macroalgae and 14 fungi were present
and, in general, these contribute toward formation of the primary and secondary films necessary for
settlement of fouling organisms. A number of sedentary organisms, i.e. the barnacles Balanus amphitrite
hawaiiensis; Balanus amphitrite malayensis; Balanus amphitrite strutsburi; Xenobalanus globicipitis, the
Table 3. Number of publications indicating records of taxa novel to the Indian Seas.
Novel means not previously recorded in the Arabian Sea, Bay of Bengal and Andaman
Sea
Years Publications Taxa

1960–1964 3 6
1965–1969 2 3
1970–1974 25 74
1975–1979 29 41
1980–1984 9 9
1985–1989 17 21
1990–1994 6 14
1995–1999 12 21
2000–2004 9 16
Total 111 205
122 D.V. SUBBA RAO
Table 4. Major taxonomic groups and taxa recorded for the first time
from the Indian Seas
Group Number of species

Algae (macro)a 11
Amphipodaa 1
Arachinnellid 3
Ascidians 13
Cirripedia-balanida 3
Bivalves 6
Cephalopodac 2
Ciliate 1
Copepoda 7
Diatoma 1
Dinoflagellate 9
Fungia 14
Gastropoda 6
Gastrotrich 6
Coelenterataa 2
Isopodaa 6
Kinorhynch 1
Mites (Halacaridae) 1
Nematoda 6
Nudibranch 3
Oligochaete 1
Ophiuroid, Holothurian 2
Ostracoda 6
Piscesc 44
Polychaetaa 19
Entoproctaa 3
Pteropod 1
Brachyura } Shrimpc 6
Tardigrada 1
Teredinid, Pholadb 3
Tintinnida 16
Turbellaria 1
Total 205
a
Fouling organisms.
b
Wood-destroying organisms.
c
Commercially important organisms.
polyzoan Beania klugei, the entoproct Barentsia ramosa, Barentsia gracilis and the polychaetes Sigambra
tentaculata and Ficopomatus enigmaticus (Mercierella enigmatica) are important fouling organisms. Of
equal, if not greater, importance was the introduction of the isopods Limnoria insulae, Limnoria unicornis,
Limnoria platycauda, the pholad Xylophaga mexicana, and the teredinids Lyrodus massa and Lyrodus
medilobata due to their wood-destroying capabilities.
DISCUSSION
Between regions with a contiguous coastline, such as the Bay of Bengal and the Arabian Sea, natural
redistributions and range extension of specific biota are possible, as in the case of the amphipods Stenothoe
gallensis, Maera pacifica, Podocerus brasiliensis and Erichthonius brasiliensis into the Arabian Sea (including
Figure 2. Number of new taxa common to the Indian Seas and other geographical regions. Dotted lines represent major shipping
routes, solid lines the area sampled, and black arrows the spread of the biota.
the Pakistan coast) from the Bay of Bengal (Venugopalan and Wagh, 1986). But the spread of biota
between geographically distinct or separate bodies of water, e.g. between the Indian Seas and the Baltic Sea
or the Mediterranean Sea, could take place only by anthropogenic activities, such as ballast water
exchanges.
Introduction of non-indigenous species has occurred sporadically over the previous 40 years, suggesting
some species may have been introduced from time to time, but did not reach sufficient density to form self-
sustaining populations. Maritime traffic could have acted as an important vector and contributed to the
accelerated introductions of alien species into the Indian Seas since 1960, more so than voluntary and
accessory introductions by the aquaculture trade, or by migratory species such as marine mammals and sea
birds traversing the hemispheres.
On a global scale, most introductions of biota are the result of ballast water carriage and release
(Carlton, 1985, 1999). Ships are the common mode of water transport, and on any given day there are
20 109 t of ballast water being transported around the world carrying an estimated 3000 to 10 000 species
(Carlton, 1985, 1999; Stewart, 1991). This heavy marine traffic introduces a variety of alien biota, and
pathogens as well, via ballast water (Ruiz et al., 2000).
Increased maritime traffic accelerates introductions of alien biota with the potential to cause harm to
native marine life and hazards to human health. Most reports of exotic introductions of fauna and flora via
ballast discharges are associated mainly with temperate seas (Carlton, 1985; Culotta, 1992; Hallegraeff and
Bolch, 1992; Carlton and Geller, 1993; Subba Rao et al., 1994; Lavoie et al., 1999; Reise et al., 1999;
Occhipinti Ambrogi, 2001), but such introductions of alien biota extend to tropical waters as well.
Physiological tolerance and acclimation of the introduced species to a wide range of conditions seems to be
the most important factor deciding their dominance over competing indigenous species, as in the case of the
invasion of Cylindrospermopsis raciborskii, a tropical freshwater bloom-forming cyanobacteria, in the mid-
latitudes (Briand et al., 2004).
124 D.V. SUBBA RAO
Introductions of exotic biota, particularly sedentary organisms, transported on the hulls of ships are also
possible. A number of studies report the probable introduction of exotic barnacles (Bhatt and Bal, 1960;
Wagh, 1974) and the fouling sphaeromatid isopod (Venugopalan and Wagh, 1987) by ships visiting the
coastal waters off Bombay. Studies of ballast water introductions into the tropical and subtropical coastal
seas, particularly to the Indian Seas, however, are largely lacking.
Prior to 1960, fishing in the area of study was artisanal and the gear deployed was selective and
inefficient compared with the industrial trawling operations that started in the 1970s. The greater frequency
of records of fish non-native to the Indian Seas from 1972 onwards coincides with the introduction of
fishing trawlers and deep-sea trawling operations by the Norwegians. Thus, better and more intensive
sampling probably resulted in finding several species of fish ‘new’ to this region. Occurrence of
Sardinella aurita in the Arabian Sea, but not in the Bay of Bengal, is attributed to migration from
its native eastern Mediterranean through the Suez Canal and the Red Sea (Dutt and Raju, 1983).
First-time presence of Parapercis alboguttata (Sreenivasan and Lazarus, 1973), Gerres macracanthus
(Venkataraman and Badrudeen, 1975), Caranx williamsi (Dutt and Shameem, 1976), Epinephelus
gauza (Dutt and Sujatha, 1984b), Richardsonichthys leucogaster (Ramanathan et al., 1974), Nemipterus
peronii (Rao and Srinivasa Rao, 1986) in these waters, mostly after a few years of trawling is significant.
In view of the excellent records on marine fish dating back to 1865 (Jones, 1971; Talwar and Kacker,
1984), these new occurrences suggest their possible introduction into the Indian Seas. Their occurrence in
large numbers, including both juveniles and adult males and females, provides putative evidence not only of
their survival, but also of their presence in a mass sufficiently critical to lead to reproduction
and establishment of self-sustaining populations. Similarly the lionfish Pterois volitans, a native to
Indo-Pacific tropical waters, which was introduced via aquarium releases or ballast water discharge into the
waters of the Atlantic coast of North America, has not only survived but also reproduces (Whitfield et al.,
2002).
Centre of xenodiversity
The East Indies Triangle, formed by the Philippines, the Malay Peninsula and New Guinea, harbours a
very rich concentration of marine species, and functions as a centre of evolutionary radiation
(Briggs, 1999); this is a view supported by this review, which also suggests that the Indo-Malaysian
region functions as a centre of xenodiversity, as evident from the 63 species common between India and
this region. This distribution is consistent with the principal shipping routes between India and other
trading partners (Figure 2). Some of the ports in the Far East and Southeast Asia are the busiest,
constituting the leading transshipment hub between the Far East and the Indian Seas (Port Industry
Statistics, 2004). According to the American Association of Port Authorities, of the 10 busiest ports in the
world by volume of the goods or containers or TEUs (twenty-foot equivalent units), more than five are
from this region. Unlike the Baltic Sea, which functions as a bridgehead for non-indigenous species
(Lepp.akoski et al., 2002) due to its temperature range of 18C to 168C and a salinity of 3 to 20 PSU, the
Indo-Malaysian region with stable hydrological conditions probably serves as a centre for the unintentional
introductions of biota via the maritime traffic from the Far East, the Phillippines, China, Japan, Australia
and New Zealand. The majority of the novel records from the Indian Seas report species common to these
waters.
Only a few species were common to the Indian Seas and South American waters (see Appendix), namely
the nematode Dolicholaimus benepapillosus (Chandrasekhara Rao, 1974) from Chile, the tintinnid
Helicostomella longa (Krishnamurthy et al., 1979) from Peru and the fish Seriola lalandi (Shameen and
Dutt, 1986) from Uruguay. The limited maritime trade activity between India and these countries explains
the paucity of new records of biota originating from the South American Pacific.
Pathogens
The potential consequences of the introduction of microbes, such as the pathogen Vibrio cholerae or
phytoplankton that cause toxigenic episodes, are of considerable concern. Some of the marine
phytoplankton and eukaryotic organisms, such as copepods, present in ballast water are believed to act
as vectors for pathogens such as V. cholerae 01, biotype El Tor associated with cholera and Cyclospora-like
bodies (CLBs) associated with diarrhoea (Epstein et al., 1994) and to have caused cholera in non-cholerae
epidemic areas such as the Peruvian coast (Epstein et al., 1994; Colwell and Huq, 2001). V. cholerae 01
and a novel serotype 0139, not previously reported in US waters, were present in 93% of foreign
ships visiting Chesapeake Bay, and the pathogens occurred 100 times more in the water than in the
zooplankton (580 mm diameter) samples (Ruiz et al., 2000). That these pathogens thrive in large
numbers in the industrially polluted iron-rich South African waters (Patel et al., 2004) and in freshwater
and marine habitats of the Chesapeake Bay and the northern Ionian Sea (Cavallo and Stabili, 2004)
without human contact is a matter of concern for epidemiologists. In fish and clams harvested off
Mangalore on the west coast of India, Karunasagar et al. (1990a) have recorded the occurrence of vibrios
pathogenic to humans, e.g. Vibrio parahaemolyticus, non-01 V. cholerae, Vibrio vulnificus and Vibrio
mimicus. On the east coast of India the shrimp aquaculture industry was worth US $100 million (equivalent
to 82 850 t yr 1 production) in 1994–1995 and was projected to generate revenue of US $300 million by
about 2000. The abrupt and total elimination of shrimp stocks (Subba Rao, 2002) was attributed to white
and brown spot diseases caused by viral, fungal and bacterial pathogens that may have originated from
ballast discharges.
Toxigenic blooms
The global spread of toxigenic (diarrhetic shellfish poisoning (DSP) and paralytic shellfish poisoning (PSP))
algal blooms due to inadvertent transoceanic and interoceanic introductions of harmful algae through
ships’ ballast discharges has been well documented (Carlton, 1985; Hedgpeth, 1993). On the west coast of
India, off Mangalore, the sporadic occurrences of PSP and DSP seem to have been caused by Dinophysis sp.
and Prorocentrum sp. blooms (Karunasagar et al., 1989). Whether any linkage exists between the
qualitative composition, abundance, viability of the ballast water biota and their potential for bloom
formation in these coastal waters remains to be established. On the west coast of India, blooms of
Alexandrium spp. have been associated with the PSP (Karunasagar et al., 1990b). The mortality of a
number of fish was associated with red water caused by Gymnodinium nagasakiense off the west coast of
India during September–October 1989 (Karunasagar and Karunasagar, 1992). That these episodes on the
west coast of India were one-time events and unrelated to season suggests an isolated sudden introduction
of the causative agent. In contrast, in the Southeast Asian region (Azanza and Taylor, 2001) and in the
temperate waters off Australia, Japan, the east coast of Canada and the USA (see Hallegraeff (1993)) such
episodes recur as a result of resident seed populations.
Pests and nuisance species

It is not only ballast water discharges, but also other shipping activities that have ramifications with
potentially grave consequences. The sediments that collect in the ballast tanks are known to harbour
pathogens, biota and cysts (Hallegraeff et al., 1990; Hallegraeff and Bolch, 1991, 1992). When a ship
is dry-docked for maintenance, the scrapings from the hull contain a variety of sedentary fouling
and boring organisms, that are usually washed into the sea and can result in localized colonizations.
Such local populations, in the long run, could contribute to competition of physical and biological
resources in a coastal ecosystem, particularly in harbours. For example, on the east coast of
India, in Visakhapatnam harbour, the black striped mussel Mytilopsis sallei, native to Atlantic
126 D.V. SUBBA RAO
waters, a congener of the zebra mussel Dressina polymorpha, was probably introduced during late 1960s
through fouled ships’ bottoms, initially occurring in relatively small numbers (Rao and Rao, 1975)
but now present in massive quantities (Rao et al., 1989). M. sallei, a dioecious, opportunistic r-strategist
with a high fecundity, settled readily in the 1980s in Victoria Harbour, Hong Kong (Morton, 1989)
and became a pest in the Pacific (Taiwan, Japan, Australia and Hong Kong). In the Indian coastal seas,
this bivalve tolerates a wide range of environmental conditions, such as 108 mg L 1 petroleum hydrocarbons
(Mohan and Prakash, 1998) and 8.364 mg Zn L 1 (Devi, 1995). Additionally, it tolerates 0–50 ppt salinity
(Raju et al., 1975), and thrives in the presence of copper and tributyltin (Karande et al., 1993). As such, this
bivalve, which has no natural predator in Indian coastal waters, attained nuisance status in Visakhapatnam
harbour resulting in an annual biomass build up of 100 kg m 2 (Rao et al., 1989). It has also spread into
Bombay waters (Karande et al., 1993), displacing much of the native fauna, and may eventually cause
drastic changes to the ecosystem. It has the potential to cause devastation in much of the Indian coastal
waters by altering the trophodynamics and a loss in native biodiversity. Alien invaders that have no natural
predators (Wolfe, 2002) or pathogens (Mitchell and Power, 2003) and seem to succeed better in the
introduced range than in their native range can attain high population densities and larger body sizes.
Furthermore, reduced levels of parasitization, as was reviewed in 26 species of molluscs, crustaceans, fish,
birds, mammals, amphibians and reptiles (Torchin et al., 2003), have similar effects.
Williamson and Fitter (1996) devised the 10:1 rule, derived from data for introduced species in Britain,
which states that, of the introduced species in a region, 10% become established, of which 10% become
pests. Of the 205 species that have been introduced into the Indian Seas, 30 taxa seem to have become
established. These are: the red algae Jania ungulate f. Brevior, Dasya sinicola Vibrio abyssicola; brown alga
Hydroclathrus tenuis; green alga Neomeris vanbosseae; polychaete Ficopomatus enigmaticus; ostracod
Sclerochilus contortus; entoproct Barentsia ramose, Bankia gracilis; isopods Limnoria insulae, Limnoria
unicornis, Limnoria platycauda; mite Atelopssalis pacific; bivalves M. sallei, Ctenoides ferescabra,
Lithophaga bisulcata, Mytilus viridis, Musculus strigatus; teredinid Lyrodus massa; cephalopod Doryteuthis
sibogae; ascidians Eudistoma constrictum, Eudistoma laysani, Eudistoma toealensis, Eudistoma ovatum,
Phallusia nigra; Botrylloides magnicoecum; and the fish Gerres macrocanthus, Decapterus macarellus,
Caranx williamsi, Nemipterus peronii, Leiognathus elongate. Of these organisms, the majority are benthic
and encrusting. Of the established taxa, only two, M. sallei and F. enigmaticus, seem to have become pests,
as evident by their massive colonization in Indian coastal waters. Boudouresque and Verlaque (2002)
described a similar relationship from data on 85 macrophytes introduced into the Mediterranean Sea.
Biofouling and wood borers

The presence of various epibionts, i.e. four balanids, two coelenterates, 19 polychaetes, three polyzoa and
13 tunicates (see Appendix), suggests they could have been introduced as adherents to ships’ hulls or some
other oceanic debris. Barnes (2002) estimated that rubbish of human origin in the sea has doubled the
propagation of bryozoans, barnacles, polychaete worms, hydroids and molluscs in the subtropics and
tripled it in the high (>508) latitudes. Besides dispersing epibionts such as hydrozoans, cyanobacteria and
macroalgae, drifting plastic debris has been suggested as a potential vector for transporting harmful algal
bloom species such as Ostreopsis sp., Coolia sp. and Alexandrium taylori along the Catalan coast (Maso!
et al., 2003). Prevention of biofouling and protection of wood from borers are of particular interest to
maritime activities. Estimated annual costs in India amount to US $400 000 for cleaning offshore oil
platforms, about US $28 million to clean the merchant navy ships, US $117 million due to corrosion (see
Rao and Griffiths (1998)) and about US $5–6 million resulting from biodeterioration of wooden fishing
craft (Santhakumaran, 1988). Preservation and protection of wood used in fishing craft and construction of
jetties, pontoons, etc. has been one of the primary goals of The Wood Preservation Institute, Dehradun.
Bacteria, fouling diatoms and algae contribute sequentially to the formation of the primary film and
secondary film necessary for the settlement of the foulers and woodborers. One pholad and two teredinids,
exotic to Indian Seas, that cause destruction of submerged wooden structures have been added in the last 20
years. Also of concern is the occurrence of the alien fouling polychaete F. enigmaticus in Visakhapatnam
harbour (Chandramohan and Aruna, 1994). Similarly, F. enigmaticus has been introduced to Europe,
where it seems to out-compete native biota and has colonized the delta area of the southwest Netherlands
(Vaas, 1975) and Emden Harbour, Germany (Kuehl, 1977), where discharges of heated effluents from
power plants have resulted in localized elevated temperatures. In the Indian coastal seas, other sedentary
organisms recently introduced include several colonial ascidians (see Renganathan (1983,1984), Meenakshi
(1998) and Meenakshi and Venugopal (2000); Appendix). And the spionid Eudistoma sp. is now established
and breeds year round (Renganathan, 1983). In the long run, these fouling ascidians could be pests similar
to M. sallei. Harbours are more vulnerable to the introduction of alien species because many ships visit
them and harbour waters are usually nutrient rich and support a good growth of plankton, thus
creating ideal nurseries (Diesel et al., 2000); the pilings and pontoons provide novel habitats for epibionts
(Connell, 2001).
Mitigation and research needs

Introduction of alien biota has many ramifications, and some of these include deleterious effects on
biodiversity, severe economic losses, and health risks. It is in the harbours that measures and development
of more effective barriers could be implemented because of limited exchange of harbour waters with the
open sea. Since 1947 (post-independence) India has been advancing industrially and with it there has been
increasing sea trade, such that it was even necessary to construct a new port, Paradip, to meet shipping
demands. Several thousands of transoceanic and interoceanic vessels call at Kolkata, Haldia, Paradip,
Visakhapatnam, Chennai, Tuticorin, Cochin, New Mangalore Port Trust, Marmugoa, Mumbai,
Jawaharlal Nehru Port Trust and Kandla each year. Additionally, there are 165 intermediate and minor
ports; and under the ‘Sagarmala’ project the Government of India proposes to modernize at least 32 of
these. There are well-used shipping routes between India and North America, west, central and east
European countries, Southeast Asia, Australia, New Zealand, West Asia and Middle East countries. At
Mumbai, a total of 3614 cargo and non-cargo vessels arrived during 2000–2001, and the total cargo
handled was 23.40 106 t whereas the other major ports, Visakhapatnam handled 32.20 106 t, Chennai
31.20 106 t, Kandla 29.59 106 t, Paradip 14.96 106 t and Kolkatta 10.20 106 t. The cumulative cargo
handled by major Indian ports was 281 106 t during 2000–2001 and it is predicted to reach 650 106 t by
2006.
Ignoring problems caused by ballast water introductions could pose a threat to the coastal bays, estuaries
and mariculture sites that sustain renewable resources and impair understanding of the biodiversity and
functioning of the ecosystem. Lack of strict regulations of ballast water discharges will exacerbate the
establishment and impact of a potentially unending tide of invaders. For the Indian Seas, long-term
institutionalized multidisciplinary research on ballast water invaders is recommended to assess the invaders,
their survival, growth and establishment of a critical mass necessary for establishment in the new domain.
Successful application of any measures to mitigate impact of ballast water introductions of alien species
is dependent on two factors: first, on the appreciation of the indigenous fauna and flora and their
functioning; second, on scientific knowledge on the quantity of introduced species, frequency and timing of
their introduction, their chances of survival in an alien environment, breeding habits, and their impact on
the local biota. Systematic studies along these lines that are focused on the major Indian harbours are
required.
The Australian Ballast Water Management Council (ABWMAC) developed a comprehensive
programme consistent with the guidelines of the International Maritime Organization (IMO), which merit
application while developing ballast water research in the Indian Seas. Usage of ballast water in oceanic
128 D.V. SUBBA RAO
ships is a century-old practice, unlike measures of mitigation and management of ballast water
introductions. Isolation of ballast water using onshore reception facilities would be effective but delay
ship sailings. A programme aimed at maritime awareness is recommended for mariners so that ballast water
exchanges and protocols will be implemented. Furthermore, ABWMAC recommends emptying and
subsequently refilling 70–90% of the ballast water outside the EEZ, which costs about 2.1 per tonne of
ballast water. Filtration through a 5 mm prescreen followed by a 25 mm screen is selectively effective, but it
does not exclude organisms 525 mm and costs 5.8 to 8.1 cents per tonne in addition to retrofitting the ships.
Similar to the offshore exchange method, it leaves behind the sediments that may contain cysts and biota.
Heating the ballast water to about 388C is another method recommended by ABWMAC; however, this
leaves resistant bacteria, pathogens, viruses and encysted stages, particularly from the warm tropical
coastal seas, unaffected. Treatment with biocides and disinfectants, such as ozone, glutaraldehyde, peracetic
acid, and chlorine, although effective, may have long-term effects on the environment. Plasma pulse
technology, which utilizes a power supply to discharge intense shock waves, a steam bubble and UV light, is
in the developmental stage by Sparktech. As no single method in itself is totally effective, it will be necessary
to use a combination of several methods to mitigate the effects of introductions of alien biota.
ACKNOWLEDGEMENTS
I am grateful to Dr J. E. Stewart, Bedford Institute of Oceanography, for constructive criticism of several versions of
the manuscript. I am indebted to Dr John M. Baxter, Edinburgh, the Chief Editor of Aquatic Conservation. Marine and
Fisheries Ecosystems, and to two anonymous reviewers for their constructive and critical comments on the manuscript.
This analysis could not have been possible without the kind help of Professor K. Srinivasa Rao. Although retired,
during a trip to India from Dallas, Professor Rao spent several weeks, located many publications and provided insights
into the bionomics and ecology of the biota from the Indian Seas. For this kind help I remain indebted to Professor
Srinivasa Rao. Special thanks are due to Dr Ravi V. Durvasula, Yale University, who encouraged me to write this
review and made many excellent suggestions on the manuscript. I am grateful to Dr C. Annapurna and Dr K. Umadevi,
my colleagues at Andhra University, Visakhapatnam, and to Dr R. Shankar, Dr K. R. Sridhar, Dr K. Jayappa,
Mangalore University, Mangalagangothri, for sending photocopies of several obscure publications. My thanks are due
to Mr Stephen Smith for statistical advice, to Mr Arthur Cosgrove and Bala T. Durvasula for assistance with the
graphics and formatting the manuscript and to Ms Marylin Rudi, Archivist Librarian, Bedford Institute for help with
literature search.
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APPENDIX: RECORDS OF BIOTA REPORTED FOR THE FIRST TIME FROM THE INDIAN SEAS AND THEIR KNOWN DISTRIBUTION
Reference Systematic Group Species Locality Natural Distribution

Ganapati and Lakshmana Isopoda Limnoria insulae Andamans S. Pacific
Rao (1960) Limnoria unicornis Andamans Florida
Limnoria platycauda Andamans W. Atlantic
Bhatt and Bal (1960) Cirripedia } barnacle Balanus amphitrite Mumbai Malay Archipelago and Persian Gulf
hawaiiensis
Balanus amphitrite Mumbai Malay Archipelago
malayensis
Ganapati and Pholad Xylophaga mexicana Puri, Bay of Bengal Mexico

Lakshmana Rao (1961)

Subrahmanyan (1968) Dinoflagellata Ceratium setaceum Arabian Sea Atlanti, Pacific
Ceratium minutum Arabian Sea Atlantic
Subba Rao (1969) Pteropoda Cymbulia peroni Visakhapatnam Atlantic, Pacific and
Mediterranean deep waters
Subrahmanyan (1971) Dinoflagellata Diplopeltopsis minor Arabian Sea Baltic, English Channel, Brittany
coast
Peridinium bulla Arabian Sea Barents Sea
Peridinium globosum Arabian Sea African Atlantic
Peridinium trochoideum Arabian Sea European coasts
Peridinium avellana Arabian Sea Plymouth, Flemish coast
BIOTA OF THE INDIAN SEAS
Peridinium cerasus Arabian Sea North Sea, Iceland, Greenland
Talwar (1972) Pisces Oxymonacanthus longi- Vizhinjam, Arabian Sea Indo-west Pacific, Philippines,
rostris Melanesia, Polynesia
Talwar (1973) Pisces Zen itea Arabian Sea South African waters
Joglekar (1973) Pisces } Hetero- Heteromycteris normani Orissa SE Asia

mycterinae
Sreenivasan and Lazarus Pisces } Pomacentridae Chromis xanthochir Viziangam Indo-Malay, Muscat
(1973) Parapercis alboguttata
Subba Rao (1973) Dinoflagellate Citharistis apsteinii Bay of Bengal Tropical Atlantic, Florida Straits,
Coral Sea, Solomon Islands,
Galapagos
Sarma and Ganapati (1973) Bivalve (epibiont on Musculus strigatus Visakhapatnam, Madagascar and Sumatra Island

137
algae) Bay of Bengal continued over

APPENDIX: continued 138
Kuthalingam et al. (1973) Pisces } Veliferidae Velifer africanus Vizianjam East Coast of Africa
Raju et al. (1974) Coelenterata } anemone Eugymnanthea inquilina Bay of Bengal Italian Seas, Puerto Rico, Japan
Wagh (1974) Cirripedia } barnacle Balanus amphitrite var. Mumbai West coast of Africa, Philippines,
stutsburi Melanesia, Polynesia
Ramanathan et al. (1974) Pisces Richardsonichthys Porto Novo SE Asia, New Guinea, China,
leucogaster Bay of Bengal Australia
Zanzibar, Muscat, Seychelles
Menon and Talwar (1974) Pisces Sardinella leiogaster Bay of Bengal Indo-Pacific

Talwar and Sathiaravan (1974) Pisces Epinnula orientalis Bay of Bengal Japan, Philippines, Gulf of Mexico
Chandrika (1974) Tunicata Kowalevskia oceanica Quilon Atlantic, Mediterranean
Chandrasekhara Rao (1974) Ciliata Epiclintes ambiguus Visakhapatnam, Baltic, Cape Cod
Bay of Bengal
Coelenterata } hydrozoa Halammohydra octopop- Baltic, North Sea, Atlantic,

dites Mediterranean
D.V. SUBBA RAO
Turbellaria Gyratrix hermaphroditus Baltic, North Sea, Atlantic,

Mediterranean, Black Sea, Florida,
Brazil
Kinorhyncha Catera styx Brazil, Angora
Gastrotricha Acanthodasys aculeatus Baltic, North Sea, Atlantic

Macrodasys caudatus Mediterranean, Florida
Urodasys viviparus Mediterranean, Florida
Thaumastoderma heideri Baltic, North Sea, Atlantic
Aspidiophorus marinus Baltic, North Sea, Atlantic
Xenotrichula velox Baltic, Atlantic, Mediterranean
Nematode Dolicholaimus Pacific, Chile, Baltic, North Sea,

benepapillosus Atlantic, Mediterranean
Oncholaimus brachycercus Pacific, Baltic, North Sea, Atlantic
Monoposthia costata Pacific, Baltic, North Sea, Atlantic
Synonchium obtusum Pacific, Atlantic, Mediterranean
Spilophorella paradoxa Pacific, Atlantic, Mediterranean
Monohytera parva Pacific, Atlantic, North Sea,

Mediterranean
Arachinnellida Nerilla antennata Pacific, Atlantic, Mediterranean
Nerillidium mediterraneum Atlantic, Mediterranean
Trilobodrilus nipponicus Pacific, Japan
Polychaete Eusyllis homocirrata Atlantic
Petita amphopthalma Atlantic, Mediterranean
Oligochaete Michaelsena subterranea Baltic, North Sea, Atlantic,
Mediterranean
Ostracod Microcythere subterranea Baltic, North Sea, Atlantic,
Mediterranean
Copepod Arenostella germanica Pacific, Brazil, Baltic, North Sea,
Atlantic, Mediterranean
Arenopontia subterranea Baltic, North Sea, Atlantic,
Mediterranean, Mozambique
Psammopsyllus opercula- Brazil, Ghana, Senegal

tus
Isopod Angeliera phreaticola Mediterranean, Madagascar
Tardigrada Stygarctus bradypus Atlantic, Baltic, North Sea
Nudibranchiata Pseudovermis salaman- Brazil
drops
Surya Rao (1974) Amphipoda Maera hirondellei Arabian Sea Azores, Ile Djerba, Algiers,
Mediterranean
Satyanarayana Rao Polyzoa Beania klugei Visakhapatnam, Tropical west Africa

and Ganapati (1974) Bay of Bengal
Ganapati and Decapoda Atanus indicus Visakhapatnam, Indo-Pacific

Kanakayya Sastri (1974) (shrimp) Bay of Bengal
Sarma and Ganapati (1974) Bivalvia Musculus strigatus Visakhapatnam Madagascar, Sumatra
Gastropoda Volvotella pyriformis Bay of Bengal
Cylindrobulla pusilla
Oxynoe viridis W. Australia, Mediterranean, Canary
Islands
Achari (1974) Polychaete Phragmatopoma sp. Cannanore, Arabian Sea Tropical Atlantic, Subtropical
Eastern Pacific, Hawaiian Islands
Patel and Mahyavanshi (1974) Brachyura Callianassa tyrrhena Gulf of Kutch Mediterranean, Adriatic, Greece
Rao KV et al. (1974) Nudibranchiata Asteronoyus caespitosus Laccadives Indian, Pacific

Phyllidia varicosa Indian, Pacific
Rao TA (1974) Pisces } Scianidae Atrobucca nibe Visakhapatnam Indo-China, Japan, South Africa

139
continued over
Sreenivasan (1974) Pisces } Carangidae Alepes macrurs Viziangam Sumatra, Java, Celebes, Philippines
Carangoides Red Sea, east and west coasts of
coeruleopinnatus Africa, Philippines, Celebes,
Formosa, Japan
Hartman (1974) Polychaeta Eupanthalis kinbergi Northern Indian Ocean North Atlantic, Mediterranean
Eupanthalis nigromaculata Philippines, East Indies
Eupolyodontes sumatranus Sumatra
Pterocirrus macroceros Gulf of Oman, Mediterranean
Gyptis arenicola Naples
Eurysyllis tuberculata Mediterranean

Ceratoneries hircinicola Mediterranean, Malagas, Red Sea
Nereis filicaudata Somaliland, Bahama Islands
Drilonereis monroi South Africa, South Atlantic
Prionospio malmgreni Mediterranean, Atlantic
Heterospio longissima Northwest Atlantic
Rajagopalan et al. (1975) Pisces } Nemepteridae Nemipterus delagoae SW Arabian Sea South Africa
Bhuti et al. (1975) Decapoda (shrimp) Alpheus splendidus Arabian Sea Gulf of California, Mexico
Khan (1975) Pisces } Molidae Mola mola Bombay Atlantic, Japan, Australia, Hawaii
D.V. SUBBA RAO
Achari (1975) Polychaeta Sigambra tentaculata Viziangam New England, California

Chesapeake Bay, Gulf of Mexico
Krishnamurthy and Tintinnidae Favella philippinensis Porto Novo Portuguese Guinea, Japan, Gulf of
Santhanam (1975) Metacyclis brevis Siam NE Atlantic
Eutintinnus tenuis
Rhabdonella sp.
Venkataraman and Pisces (silverbidde) Gerres macracanthus Palk Bay Formosa, Indonesia
Badrudeen (1975)
Jayaprakash (1975) Pisces } Scianidae Nibea chui Bombay W Pacific
Satyanarayana Rao et al. Gastropoda Distorsio reticulata Palk Bay Japan

(1976)
Thomas (1976) Decapod (shrimp) Alpheus rapax Palk bay Red Sea, East Africa,
Mergui Archipelago

Sarvesan (1976) Cephalopoda } cuttlefish Sepia trygonina Tuticorin Red Sea and Gulf of Aden
Lal Mohan and Gandhi (1976) Pisces Ypsiscarus oedema Gulf of Mannar Celebes Sea, Philippines, Japan,
Tahiti
Dutt and Shameem (1976) Pisces } Carangidae Cranx williamsi Visakhapatnam, Kenya
Kakinada
Satyanarayana Rao et al. Bivalve Mytilopsis sallei Visakhapatnam Caribbean, Hong Kong, Central
(1976) America, Mexico, Panama
Sreenivasan (1976) Pisces } Carangidae Decapterus macarellus Vizhinjam, Arabian Sea Australian seas, Atlantic, Pacific
Appukuttan (1976) Bivalve Lithophaga bisulcata Gulf of Mannar East and west coasts of America

Thompson and Meiyappan Copepoda Monstrilla turgida Arabian Sea, Minicoy Atlantic, Mediterranean, Indo-Pacific
(1977)
Saraladevi (1977) Copepod Haloptilus paralongicirrus Arabian Sea and Northern Atlantic
Haloptilus bulliceps Bay of Bengal
Raman and Ganapati (1977) Polychaete Scolaricia capensis Visakhapatnam South Africa
Peter (1977) Polychaete Plotohelmis sumatraensis Arabian Sea, Sumatra

Bay of Bengal
Devarajan and Brachyura Ixa inermis Hong Kong, China

Girijavallabhan (1977)
Thomas (1977) Isopoda Epipenaeon ingens Palk Bay, Red Sea

Bay of Bengal
Patel (1977) Diatom Biddulphia punctata Antarctic
Patel (1978) Bivalve } Lamellibranch Ctenoides ferescabra Gulf of Kutch Low Isles, Great Barrier Reef
Murty (1978) Pisces } Nemepteridae Nemeipterus mesoprion Kakinada Sumatra, Singapore
Krishnamurthy et al. (1978) Tintinnidae Rhabdonella spiralis Porto Novo Somali Current
Krishnamurthy et al. (1979) Tintinnidae Tintinnopsis butchilii Porto Novo, Atlantic, Japan
Bay of Bengal
Tintinnopsis uruguayensis Porto Novo, Brazil, Mandacaru River
Bay of Bengal

141
continued over
Coxliella annulata Porto Novo, Bay of Atlantic
Bengal
Coxliella ampla Porto Novo, Bay of Atlantic, Japan
Bengal
Metacylis jorgensenii Porto Novo, Bay of Portugal, Atlantic, Japan
Bengal
Favella brevis Porto Novo, Bay of Atlantic
Bengal
Tintinnopsis lohmani Atlantic, Japan
Tintinnopsis bermudensis Bermuda
Tintinnopsis uruguayensis Mandacaru River, Brazil
Tintinnopsis beroidea Atlantic, Japan

Helicostomella longa Peru, Japan
Siva Rama Sarma and Polychaeta Pisione oestedii Visakhapatnam South Africa
Chandra Mohan (1979)
Rama Rao (1999) Pisces } Scorpionidae Pteroidichthys amboinensis Madras Celebes, Vietnam
Untawale et al. (1980) Alga Monostroma oxyspermum West coast of India Galicia (Spain), NE Atlantic, Japan,
NW Pacific
Renganathan (1981) Tunicata } Ascidian Styela bicolor Bay of Bengal Gulf of Siam, Java, N. Australia,
D.V. SUBBA RAO
Banda Sea, Philippines
James (1982) Ophiurid Ophiocoma anaglyptica Lakshdweep Canton, S. Pacific Islands
Dutt and Raju (1983) Pisces } Clupeidae Sardinella aurita Quilon Atlantic, Mediterranean, W. Pacific
Santhakumaran (1983) Teredinid Lyrodus massa Goa Yemen, Puerto Rico, Eritrea, Kenya,
South Africa, east coast of
Madagascar, Singapore
Dutt and Sujatha (1984a) Pisces } Sillaginidae Sillago intermedius Visakhapatnam Thailand
Dutt and Sujatha (1984b) Pisces } Serranid Epinephelus guaza Visakhapatnam Eastern Atlantic, Brazil,
Mediterranean
Renganathan and Monniot Tunicata } Ascidian Aplidium multiplicatum Bay of Bengal Japan
(1984)

Silas et al. (1985a,b) Cephalopoda } squid Doryteuthis sibogae Vizhinjam, Madras, Indonesia, Formosa, Taiwan
Trivandrum
Renganathan and Tunicata } Ascidian Botrylloides magnicoecum Tuticorin South Africa, Bermuda,
Krishnaswamy (1985) New Zealand
Kumari et al. (1985) Pisces Polynemus sheridani Bay of Bengal Northern Territory of Australia
Silas and Selvaraj (1985) Pisces (ray) Dasyatis centroura Off Quilon Atlantic, Adriatic, Mediterranean
Shameem and Dutt (1986) Pisces Carangoides dinema Visakhapatnam South Africa, Tanzania, Papua New
Guinea, Tanga Islands

Seriola rivoliana Mediterranean, Azores, Devon (UK)
Uruguay
Seriola lalandi NSW (Australia), New Zealand,

eastern Pacific, Japan, Uruguay,
False Bay, South Africa
Seriola dumerili Japan, Gulf of Mexico, Mediterra-

nean, NW Sicily, Adriatic
Rao and Srinivasa Rao (1986) Pisces } Nemipteridae Nemipterus peroni Tuticorin S. China Sea
Rama Murty and Srinivasa Pisces (flat fish) Cyanoglossus lachneri Paradip, Visakhapatnam East Africa, Arabia, Red Sea,
Rao (1986) Seychelles
James (1986) Holothuridae Phyrella fragilis Port Blair Japan
Santhakumaran (1986) Teredinid Lyrodus medilobata Gujarat Indo-Pacific, Hawaii, Marshall

Islands, New Caledonia,
New Zealand, Australia
Sujatha and Dutt (1986) Pisces } Serranid Epinephelus hata Visakhapatnam Japan
Kasinathan et al. (1986) Gastropoda Marginella ventricosa Porto Novo Indonesia, Malay, Japan
Venugopalan and Wagh Isopoda Cilicaea latreilli Bombay Indonesia, Philippines, South Africa,
(1987) Red Sea, Australia
Jayachandran and Joseph Decapoda (prawn) Macrobrachium aemulum SW Arabian Sea New Caledonia
(1988)

143
continued over
Ravikumar and Fungi Corollospora intermedia Vellar estuary Helgoland (Germany)
Purushothaman (1988)
Devadoss (1988) Pisces (fantail ray) Taeniura melanospila Madras E. Africa } Batavia, East Indies
Satyanarayana Rao et al. Entoprocta Barentsia gracilis Visakhapatnam Italy, Senegal, N. Pacific
(1988)
Barentsia ramosa Pacific, California, Belgium
Ambiye et al. (1989) Alga } Chlorophyte Neomeris vanbosseae Laccadives Kenya
Chinnaraj and Untawale Fungi } Ascomycetous Rhizophila marina North Sumatra, Seychelles

(1992) Trematosphaeria
striatipora
Lineolata rhizophorae Maldives Mauritius, Japan
Caryosporella rhizophorae Belize
Passeriniella savoryellopsis
Hypoxylon oceanicum Hong Kong, Malaysia
Belizeana tuberculata Belize
Swampomyces armeniacus N. Queensland (Australia)
Hydronectria tethys var. Arabian Sea SE Asia, Mexico, Fiji, USA, Belize
glabra
D.V. SUBBA RAO
Sarma and Chatterjee (1993) Halacaridea (mites) Atelopsalis pacifica Port Blair Mactan Island, Philippines
Ali-Khan (1993) Copepoda Benthomisophria palliata Arabian Sea Azores
Gopakumar et al. (1993) Pisces } Mackerel Scomber australasicus Vizhinjam, Kerala New South Wales, New Zealand,
Japan, Taiwan, Red Sea
Ingole et al. (1994) Bivalve (mangrove clam) Geloina erosa Mandovi estuary Indo-Pacific, Malay
Chandramohan and Aruna Serpulid Ficopomatus enigmaticus Indian waters Australia, Baltic
(1994)
Srinivasarengan et al. (1995) Pisces } Leiognathidae Leiognathus elongatus Madras W. central Pacific, West Indies
Sujatha (1995) Pisces Myrophis lepturus Visakhapatnam China

Uranoscopus bicinctus China
Uranoscopus oligolepis China
Lagocephalus guentheri China
Jagtap and Untawale (1996) Brown alga Hydroclathrus tenuis Kutch China

Sundararajan et al. (1996) Alga (calcareous algae) Halimeda distorta Pamban Caroline Islands
Shameem and Rao (1997) Pisces Nemipterus marginatus Visakhapatnam Indo-Australian Archipelago
Shameem and Rao (1998) Pisces Johnieops distinctus Visakhapatnam China, Japan, Taiwan Strait
Meenakshi (1998) Tunicata } Ascidian Phallusia nigra Tuticorin Bermuda, Brazil, Red Sea, Gulf of
Eusynstyela tincta Aden
Meenakshi and Renganathan Tunicata } Ascidian Ascidia sydneiensis Tuticorin Australia, Indonesia, Japan, Africa,
(1998) Atlantic Ocean
Annapurna and Sarma (1999) Ostracoda Loxoconchella honolulenis Visakhapatnam Easter Island, Hawai, New Guinea,

East Africa, Persian Gulf,
Samoa, New Caledonia, Western
Australia
Annapurna (1999) Ostracoda Loxoconcha sinensis Visakhapatnam Hong Kong, Sea of Japan, Booby
Island (Australia), Indo-Pacific
Loxoconcha sculpta Booby Island (Australia)
Loxoconcha illjeborgii Indo-Pacific waters
Rao UM (1999) Red algae Jania ungulata Visakhapatnam Brazil, Mauritius, Hong Kong, Japan
Dasya sinicola v. abyssi-
cola
Prassannarai et al. (1999) Fungi Arenariomyces majusculus Mangalore Red Sea, Philippines
Didymosphaeria ligno-
maris
Nimbospora effusa Australia
Tewari et al. (2000) Bivalve mussel Mytilus viridis Gujarat, Kerala, Tropical and subtropical Pacific
Karnataka
Meenakshi (2000) Ascidian Trididemnum cerebriforme Tuticorin South Africa, Australia, New Zea-
land, Philippines, Japan, Hawaii, Fiji
Meenakshi and Venugopal Ascidian Ecteinascidia sluiteri Bay of Bengal Australia, Palau Islands
(2000)
Prasannarai and Fungi Amylocarpus Mangalore USA, Hawaii

Sridhar (2001) encephaloides
Sunilkumar (2001) Gastropod (mud snail) Hydrobia sp. Cochin, Arabian Sea Indo-Pacific, South Africa, Malaysia

145
continued over
146
APPENDIX: continued
Kar and Chakraborty (2001) Pisces } Synoidae Saurida wanesio West Bengal Japan
Annapurna (2001) Ostracoda Sclorochilus contortus Visakhapatnam Pan-Arctic, Scandinavia, Greenland

Rao UM (2002) Macroalgae Enteromorpha salina Mandapam, Gulf of Florida, Louisiana, Bahamas
Mannar
Gelidiopsis gracilis Gulf of Mannar Barbados, French Guyana
Tenaciophyllum lobatum Idinthakarai coast Mauritius, Tanzania
Vadalia fimbriata Gulf of Mannar Danish West Indies, Mauritius,
Tanzania
Laurencia elata Pamban, Gulf of South Australia
Mannar
Meenakshi (2002) Ascidian Eudistoma constrictum Mandapam Australia

Eudistoma laysani Australia, Palau Islands, Indonesia,
D.V. SUBBA RAO
Philippines, Hawaii
Eudistoma ovatum Australia, Palau and Gilbert Islands,
Indonesia
Eudistoma toealensis Kei Islands, Toeal, Indo-Pacific

Biota of India - Subba Rao

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AQUATIC CONSERVATION: MARINE AND FRESHWATER ECOSYSTEMS

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Comprehensive review of the records of the biota of the

D.V. SUBBA RAO*

The geographic area

Baseline data on biota of the Indian Seas

Publications on distribution of marine fauna and ﬂora

Category of study Up to 1960 1961 to 2004

Collation of details on non-native taxa

Years Publications Taxa

Group Number of species

Pests and nuisance species

Biofouling and wood borers

Mitigation and research needs

Reference Systematic Group Species Locality Natural Distribution

Ganapati and Pholad Xylophaga mexicana Puri, Bay of Bengal Mexico

Copyright # 2005 John Wiley & Sons, Ltd.

Peridinium cerasus Arabian Sea North Sea, Iceland, Greenland

Joglekar (1973) Pisces } Hetero- Heteromycteris normani Orissa SE Asia

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

algae) Bay of Bengal continued over

Copyright # 2005 John Wiley & Sons, Ltd.

Chandrika (1974) Tunicata Kowalevskia oceanica Quilon Atlantic, Mediterranean

Coelenterata } hydrozoa Halammohydra octopop- Baltic, North Sea, Atlantic,

Turbellaria Gyratrix hermaphroditus Baltic, North Sea, Atlantic,

Gastrotricha Acanthodasys aculeatus Baltic, North Sea, Atlantic

Nematode Dolicholaimus Paciﬁc, Chile, Baltic, North Sea,

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Copyright # 2005 John Wiley & Sons, Ltd.

Satyanarayana Rao Polyzoa Beania klugei Visakhapatnam, Tropical west Africa

Ganapati and Decapoda Atanus indicus Visakhapatnam, Indo-Paciﬁc

Kanakayya Sastri (1974) (shrimp) Bay of Bengal

Rao KV et al. (1974) Nudibranchiata Asteronoyus caespitosus Laccadives Indian, Paciﬁc

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Copyright # 2005 John Wiley & Sons, Ltd.

Achari (1975) Polychaeta Sigambra tentaculata Viziangam New England, California

Jayaprakash (1975) Pisces } Scianidae Nibea chui Bombay W Paciﬁc

Satyanarayana Rao et al. Gastropoda Distorsio reticulata Palk Bay Japan

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Copyright # 2005 John Wiley & Sons, Ltd.

Peter (1977) Polychaete Plotohelmis sumatraensis Arabian Sea, Sumatra

Devarajan and Brachyura Ixa inermis Hong Kong, China

Thomas (1977) Isopoda Epipenaeon ingens Palk Bay, Red Sea

Patel (1977) Diatom Biddulphia punctata Antarctic

Murty (1978) Pisces } Nemepteridae Nemeipterus mesoprion Kakinada Sumatra, Singapore

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Copyright # 2005 John Wiley & Sons, Ltd.

Banda Sea, Philippines

James (1982) Ophiurid Ophiocoma anaglyptica Lakshdweep Canton, S. Paciﬁc Islands

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Copyright # 2005 John Wiley & Sons, Ltd.

Seriola lalandi NSW (Australia), New Zealand,

Seriola dumerili Japan, Gulf of Mexico, Mediterra-

Rao (1986) Seychelles

James (1986) Holothuridae Phyrella fragilis Port Blair Japan

Santhakumaran (1986) Teredinid Lyrodus medilobata Gujarat Indo-Paciﬁc, Hawaii, Marshall

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Ambiye et al. (1989) Alga } Chlorophyte Neomeris vanbosseae Laccadives Kenya

Copyright # 2005 John Wiley & Sons, Ltd.

Ali-Khan (1993) Copepoda Benthomisophria palliata Arabian Sea Azores

Sujatha (1995) Pisces Myrophis lepturus Visakhapatnam China

Aquatic Conserv: Mar. Freshw. Ecosyst. 15: 117–146 (2005)

Copyright # 2005 John Wiley & Sons, Ltd.