Compagno (1990a) proposed a series of habitus types or ecomorphotypes, characteristic patterns of morphology, habitat, and activity that can be used to classify cartilaginous fishes and to subdivide them ecologically.
The very varied ecomorphotypes of sharksand chimaeroids in the Region (Table 6) indicate the wide taxonomic and
Table 6. Ecomorphotype coding of sharks and chimaeroids in the Region, with numbers of species in each category and % of total number of species.
Ecomorphotypes Littoral cancritroph (sharks) Littoral eurytroph (sharks) Littoral (sharks)
Littoral sphyrnid (sharks) Littoral teuthotroph (sharks) Macroceanic (sharks) Macroceanic filter feeder Microceanic (sharks) Archipelagic (sharks) Tachypelagic (shark) Tachypelagic filter feeder
Code
ecological variety of sharks and chimaeroids in the area.
Of these, the variants on the primitive active littoral body form of shelf-dwelling sharks, including macropredatory (littoral eurytroph), specialist cephalopod (teuthitroph) and crustacean (cancritroph) feeders, and the hammerheads (littoral sphyrnids), are the most important. A large component of bottom-dwelling shelf sharks and chimaeras with various benthic morphotypes also occur in the area, including compressed chimaerids (chimaerobenthics), generalised bottom-dwellers (probenthics), elongated bottom-dwellers (leptobenthics), sawfish-like forms (pristobenthics, for sawsharks), and flattened angel-shark like forms (squatinobenthics). Deep-slope morphotypes among sharks include elongated eel-like forms (anguilliform bathics, for the frilled shark Chlamydoselachus anguineus), generalised deepwater 'floaters' (bathics, for many slope squaloids and deepwater lamnoids), and long-nosed deepwater forms (rhynchobathics, including the goblin shark Mitsukurina owstoni).
There are morphotypes in the Region for specialised oceanic sharks, including large macroceanic sharks (lamnoids and carcharhinids, plus the possibly macroceanic filter-feeding whale shark Rhincodon typus) and dwarf microceanic sharks (primarily dalatiid squaloids, also the crocodile shark Pseudocar charias kamoharai).
Adaptations for sustained cruising and high speed are seen in the few tachypelagic sharks in the area and worldwide, including the shortfin mako Isurus oxyrinchus and its essentially macroceanic-converted relative the longfin mako I. paucus, with the marginally (Taiwan) distributed basking shark Cetorhinus maximus a tachypelagic morphotype converted to slow but steady filter-feeding on small crustaceans. The archipelagic morphotype is only represented by the great white shark, which shows tachypelagic characteristics combined with adaptations for predation on large marine vertebrates.
Discussion
The Region has a rich tropical chondrichthyan fauna which is best known from the broad shelves, which support a wide variety of artesanal and commercial fisheries as well as small to huge fish markets in the various countries. As has been known by systematic ichthyologists since the last century (most notably from Pieter Bleeker's work in Indonesia and elsewhere in the tropical Indo-West Pacific in the last century), fish markets in the Region are excellent places for obtaining a broad sample of the local inshore and increasingly offshore and oceanic ichthyofauna, including rare and unusual species. Cartilaginous fishes are mostly caught as bycatch of other fisheries (including high-technology oceanic fisheries) driven by more fecund bony fishes and other fisheries species (Compagno 1990b).
The markedly increasing value of shark fins in general and
large species in particular during the last decade (including the fins of sharkfin guitarfishes and sawfishes) encourages development and expansion of specialist fisheries targeting large sharks in the Region for fins and for local consumption of their meat, as well as removal of fins from small bycatch sharks during processing of their carcasses for human consumption or other uses.
An important feature of tropical inshore fisheries for sharks is the bycatch of the young of large, more slow-growing species such as tiger sharks, as well as young and adults of small and medium-sized sharks. Many species of sharks use inshore waters as nursery grounds, and small to large scale fisheries in the Region impact these directly. It is suspected that some of the larger species may have declined in the Region due to increased mortality of the young from expanding coastal fisheries, as well as targeted fisheries. Targeted fisheries for large and medium-sized coastal sharks cut down recruitment of young by decreasing the number of breeding adults (Holden's model, 1974), while massive bycatch fisheries that land small sharks, as well as small-boat fishers that target small sharks or collect them as bycatch, catch the young of the larger species and cut down the recruitment of adults.
Fishing methods in the Region are traditionally varied and increasingly modern and intensive. They include poison and explosive-fishing, use of SCUBA gear, and high-technology local and international fishing vessels and gear to augment traditional fishing methodology, artisanal fisheries, and low-tech commercial fisheries (including inshore bottom trawling fisheries with locally manufactured moderate-sized boats and gear) without replacing them.
Unlike selective fisheries and markets in North American and some European countries, cartilaginous fishes are generally landed and utilised in tropical markets in the Region and in most parts of the world as part of traditional 'catch-everything fisheries' in which marginal and sometimes useless and toxic species are landed as well as species that are readily utilised for human consumption. Offshore high-technology international fisheries with long-range fleets may ditch finned sharks or bring them in, but the fins are utilised regardless. Cartilaginous fishes form less than 1%
of world fisheries landings according to FAO statistics.
Non-selective, expanding fisheries are essentially driven by increasing markets for fisheries products and sustained by more fecund and fishable species than chondrichthyans, including teleosts, cephalopods and crustaceans. Declining chondrichthyan catches in such non-targeted fisheries have little influence on the continuance of the fisheries. In such fisheries chondrichthyans and other K-selected vertebrates may be caught in the screws of an r-selected meatgrinder, so to speak, and with inadequate monitoring can disappear locally while the fisheries continue at high levels.
Market sampling can be an extremely valuable tool for estimating the biodiversity of sharks and other chondrichthyans in the Region, including determining the
relative abundance of various fisheries species over time as the burgeoning fisheries take their toll. It should be noted, however, that market sampling cannot substitute for wide-ranging faunal surveys of the Region, as the markets often reflect the activities of local fishers who can be relatively conservative and selective in optimising the best catches on well-known inshore grounds. Markets in the Region are, for the most part, not being monitored in detail for species-specific data on chondrichthyan catches, including intraspecific composition by sex, size and age class, and this is reflected by the datasets on cartilaginous fishes provided to the United Nations Food and Agriculture Organization (FAO), with most countries in the Region combining chondrichthyan catches for various species or providing separate statistics for sharks and rays. There are no statistics available for chondrichthyan catches off Vietnam, Brunei, and Cambodia.
The Sabah project is one of the first attempts in the Region to provide intensive species-specific monitoring of tropical markets that land primarily inshore elasmobranchs (as well as actively searching for freshwater elasmobranchs), but it faces problems of continuity, funding, and trained personnel to do the intensive and often laborious and unpleasant field work. What is necessary is long-term and broad-based species-specific monitoring of chondrichthyan catches, which is properly a function of fisheries agencies in cooperation with systematists and universities, but which is not occurring in most of the world. Systematics is the essential basis of biological research, including fisheries biology, yet systematists in general, and chondrichthyan systematists in particular, are relatively few and declining in numbers, while the need for systematic research including alpha taxonomy has increased markedly over the past few decades with the expanding human population and commensurate impact on terrestrial and marine environments. Systematics has become unfashionable in many universities; well-established university systematics institutions such as the Division of Systematic Biology at Stanford have disappeared, and recruitment of new systematists and employment of young postdoctoral systematists is not tracking the increased need for such researchers. There is often no incentive for the present generation of established systematic researchers to train replacements in the form of graduate students, and no incentives for graduate students to become systematists if they face an uncertain career.
There is evidence from ichthyological sampling in the 1960s by the George Vanderbilt Foundation and more recent market surveys in Thailand in 1993 and 1995, that certain groups of inshore chondrichthyans, including sawfish (Pristidae), possibly electric rays (Narcinidae and Narkidae), and eagle rays (Myliobatidae), have markedly declined in the Gulf of Thailand, while shark diversity and landings have decreased and whiptailed stingrays (Dasyatidae) and sharkfin guitarfishes (Rhinidae) are
increasingly dominating the declining catches. Declining inshore elasmobranch catches off the Philippines and Thailand as recorded by FAO fisheries statistics (FAO 1996) are worrying, as are increasingly massive and probably unsustainable catches of elasmobranchs off Indonesia; these reached 93,000 metric tonnes in 1994 (the highest in the world). The region includes several countries that have major elasmobranch landings (over 10,000t per year), including Malaysia, Indonesia, Taiwan and China, and formerly Thailand and the Philippines.
The shelf shark fauna of the Region is best known from off China and Taiwan, the Gulf of Thailand, Singapore and Philippines. Surveys in the Gulf of Thailand and off Sabah reveal a number of new records of sharks and other elasmobranchs, and suggest that even the inshore elasmobranch fauna of the Region is imperfectly known and needs more survey work to improve our knowledge of it. There is a danger that offshore benthic shelf chondrichthyan faunas may be adversely impacted as fisheries expand into areas that are poorly known by systematists and fisheries biologists.
The oceanic chondrichthyan fauna of the epipelagic zone in the Region is well-known due to extensive collecting as part of fisheries investigations, but is relatively undiverse and unspeciose as elsewhere in the world. The slope fauna is best known off Taiwan (where deepwater sharks are routinely landed in the local fisheries), southern China and the Philippines, but is poorly known elsewhere. With the rush to exploit deepwater teleosts elsewhere in the world (particularly in the North Atlantic but increasingly in the southern hemisphere), it is quite likely that deepwater fisheries will expand in the Region and will have a negative impact on a little-known chondrichthyan fauna including deep-slope sharks, skates and chimaeroids.
Acknowledgements
This paper is dedicated to my late colleague and good friend Sid Cook. Sid was an inspiration and supported some of the work presented here through his involvement in a project to produce a shark parts guide for the US Ocean Wildlife Campaign. We had good times doing qualitative surveys of elasmobranchs in numerous fish markets in Sabah, Peninsular Malaysia, Thailand and Singapore. Special thanks to Sarah Fowler of the IUCN Shark Specialist Group, for inviting and supporting my participation in this conference, Mabel Manjaji and Frances Dipper of the Sabah project, Alvin Wong of Sabah Fisheries, Merry Camhi of the US National Audubon Society and Shark Specialist Group, Tyson Roberts of the California Academy of Sciences, Peter Ng, Kelvin Lim, and Swee Hee Tan of the National University of Singapore, Kent Carpenter and Volker Niem of FAO, Anna Wong of the Sabah State Museum, Jack Randall of the Bishop Museum,
Peter Last of CSIRO Hobart, Bernard Seret of the Museum National d'Histoire Naturelle, Paris, and Supap Monkolprasit and Prachit Wongrat of Kasetsart University, Bangkok.
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